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Home , Anthony Road, Emu River (Tasmania), Forth River (Tasmania), Granville Harbour, Marlborough Highway, Megascolecidae

Distribution and Conservation Status of Tasmanian Native

Dr Tim Kingston

The Distribution and Conservation Status of Tasmanian Native Earthworms Project, Project No. 9214, was carried out with the assistance of funds made available by the Commonwealth of under the National Estate Grants Program (State Component). The Distribution and Conservation Status of Tasmanian Native Earthworms

NEGP PROGRAM 9214

Dr Tim Kingston Queen Victoria Museum Launceston

What was the background to this project?

During the period 1986 to 1990, while employed by the Tasmanian Department of Primary Industries and Fisheries, the author was involved in a research project examining the benefits of exotic earthworms in Tasmanian pasture . Surveys of earthworms on farmland revealed that wherever pasture had been established for some years through cultivation and the addition of fertilizers and lime, introduced of earthworms ol'1ly were found. On more recently established low production pasture, mostly on poor soils, and also on very recently cleared land some native earthworms were found to have survived (Kingston and Temple-Smith 1989). That particular study, together with others reported in the literature, derT}onstrated that earthworms have the potential to improve structure following compaction and other physical degradation and that pasture production may be enhanced by significant amounts through introduction of earthworms to newly established pasture (Temple-Smith et al 1993). An earlier study by the author (Kingston 1989) had also shown that under adverse conditions. in this case irrigation, trampling by grazing can severely deplete populations. These results taken together suggest that earthworm population density in cultivated soils might have the potential to act as a biological indicator of soil health.

Some preliminary observations by the author during the above studies suggested that native earthworms were widespread, diverse and abundant in undisturbed soils under native vegetation, in some cases only a short distance outside a fence line separating pasture from bushland. Although introduced earthworm species were sometimes found in bushland, their occurrence was invariably associated with soil disturbance, whether severe, such as at log loading sites and fire dams, or minor, such as beside walking tracks. These observations begged exactly the same kind of questions for native earthworms in bushland, most particularly in relation to forestry activities in native forest, as those discussed above for farmland. In considering soil health under forestry or other anthropogenic changes in native vegetation, the spotlight falls largely on the survival of native earthworm species; although the extent of incursion by introduced species may have value as an additional measure of soil disturbance. When dealing with native species, especially where there is partial or complete local clearance of native vegetation, there is the additional but vital consideration of the irnpact of this activity on the conservation status of earthworm species themselves. The likely severity of the impact on individual earthworm species will clearly be potentially greatest for species that are most limited in their geographical range as well as those confined to soils most highly prized for cultivation. Other pertinent questions in this context relate to the likely number of native earthworm species that have already been deleted by the extensive clearing of native vegetation that has occurred, and is still taking place in ; the value of bushland remnants for species' survival and the ability of native earthworms to recolonise and displace exotic species when native trees are replanted into former pasture.

Whether one's interest lies in earthworms as agents of soil remediation, as indicators of structural decline, or in the status of earthworm populations and species, there is a clear need to recognise the identity of the particular species, single or multiple, that are present in the location of interest. In 1990 the primary focus of the research of the author, now at the Queen Victoria Museum, migrated from exotic earthworms in farmland to native earthworms wherever they could be found. Specific studies of the impact of common forestry practices in Tasmanian forests were planned and some preliminary sampling conducted so that familiarity with the species could be attained. A search of the literature on Tasmanian native earthworms at that time revealed that the only recent paper dealing with the identity of Tasmanian native earthworms was a paper entitled "The Earthworms of Tasmania" by Australia's leading earthworm taxonomist, Professor Barrie Jamieson and published in 1974 (Jamieson 1974). This catalogue contains detailed descriptions and names of some 46 native earthworm species known from the State.

Attempts to identify the species in hand with the aid of this catalogue met, for the majority of species, with remarkably little success. Closer examination of the catalogue revealed that its contents were based upon collections made at a rather limited number of sites across Tasmania, that several sites were clustered around and two other localities and that collecting had been perfunctory rather than systematic at many of the sites.

Having become aware of the serious limitations in the quantity and geographical representation of the collections upon which Jamieson (1974) was founded, and the extent of the impediment to ecological studies that this represented, the author diverted his attention from the proposed studies of the responses of earthworms to disturbance, to rather more basic studies of earthworm diversity, distribution and . Necessarily, the first step was to undertake a comprehensive collecting programme of native earthworms throughout the State, just such a programme was conducted by the author and funded by the National Estate Grants Program for the two years 1992/93 and 1993/94.

2 How much was known about Tasmanian native earthworms prior to this project?

Despite a promising start in the late eighteenth century, particularly by Baldwin Spencer who described 18 species in 1895, the study of Tasmanian native earthworms has been neglected ever since, as the maxim "out of sight, out of mind" appears to have largely prevailed during the twentieth century.

Since the time of Baldwin Spencer, very few additional specimens had found their way into permanent collections until the efforts of Professor V.V. Hickman and more especially Dr John Hickman at the University of Tasmania conducted sporadic collecting, initially for use in student practical classes, then to provide material for more focussed research by John Hickman, and eventually specifically to support taxonomic studies by Professor Barrie Jamieson at the University of . By the time Professor Jamieson was ready to publish his summary of the known Tasmanian earthworm species in 1972, he had available to him approximately 400 specimens from about 65 collection locations around Tasmania. A summary of these sites (Table 1) however reveals that many of them were concentrated in a few frequently collected areas, indeed a third of them (22) were from just "Hobart", "Mt Wellington" and "". Within this material Jamieson found representation of 8 previously described species and material that was the basis for the description of 26 new species. The remaining 12 species, reported for Tasmania in the early literature, were not found among the recent col/ections available to Jamieson, further reinforcing the emerging realisation that the group was as yet poorly sampled in the field.

Before his 1974 paper appeared in print, Professor Jamieson visited Tasmania on a brief collecting trip. With companion Mr Terry Walker, he drove from Launceston to the east coast via St Marys and St Helens to Hobart, from there to Queenstown via Bothwell and and then to Somerset on the north coast via , Rosebery and Tullah. They collected a further 400 specimens from 23 locations during this 5 day journey.

Following this col/ecting trip, Professor Jamieson conducted some initial sorting and identification of the material, but the emphasis of his work had by then developed in a different direction. Professor Jamieson donated his collections to the Queen Victoria Museum in 1991. The geographical origins of the specimens are summarised in Table 2.

Preserved collections made between 1974 and 1990 were again few, notable among them being those of the Lower Gordon Scientific survey conducted in 1976 and donated to the Queen Victoria Museum by Dr John Hickman in 1991 and some pitfall collections made by Dr Bob Green at Maggs Mountain (Arm River), during ecological studies in that area during the period 1975 to 1990.

Collections of earthworms under the author's program commenced in 1990. Firstly, Dr Bob Mesibov was contracted to conduct an intensive survey of an are of limited geographical extent close to Launceston. Mount Arthur, near Lilydale.

3 TABLE 1 Collections made between 1930 and 1972 and incorporated in Jamieson 1974

n spp date locality locality detail lat. & long. collectors Feb 1892 Launceston 147-10 41-25 Simson, A 1 1 ?Jan 1893 ? ? Baldwin Spencer Collection 12 2 27-Jan-38 New Harbour 146-10 43-30 King, C.D. 1 1 19-Apr-38 Kallista Ck 146-35 42-45 Rodway, Mrs 8 1 Nov 1938 Coxs Bight 146·15 43·30 King,C.D. 1 1 19-Jan-39 Maydena Snowy Mountains 146-40 43-00 King, C.D. 3 16-Feb-39 Maydena Russel River 146-40 42-45 King, C.D. 5 3 Jan 1940 Kelly's Beach 145-55 43-20 King, C.D.? 6 3 Feb 1941 Lake St Clair 146·10 42-05 (Kershaw?) 5 2 26-Jun-47 Risdon Risdon 147-20 42-50 Hickman,V.V. 1 1 28-Jun-47 Mt Wellington 147·15 42-55 Hickman,v.V. 2 1 13-Sep-51 Mt Wellington 147-15 42-55 Hickman,V. V. 1 1 31-Mar-52 Mt Wellington 147-15 42-55 Radford, W. 1 1 26-Aug-53 Weldborough Pass 1.6 miles from eastern end of pass 147-55 41-10 Hickman, J.L. 1 1 26-Aug-53 St Helens Launceston via Scotsdale, 94/96 mile posts 146-10 42-05 Hickman, J.L. 1 1 26-Aug-53 Lilydale Mt Arthur 147-55 41-10 Hickman, J.L. 1 31-Aug-53 Lenah Valley New Town creek 147-15 42-55 Hickman, J.L. 2 2 11-Sep-53 Hobart Mt. Nelson, Sandy Bay 147-20 42-55 Hickman, J.L. 8 2 15-Sep-53 Lenah Valley track along Newtown Ck 147-15 42-55 Hickman, J.L 2 15-Sep-53 Hobart Waterworks Road 147-20 42-50 Hickman, J.L 2 1 31-Dec-53 I nterview River 2 miles inland, south of Interview River 144-55 41-35 Jackson, R 2 1 30-Nov-53 Tarraleah 146-25 42-20 Hickman, V.V. 3 2 13-Jan-54 Mt Wellington 147·15 42·55 Hickman, J.L. 6 3 04·Mar-54 Mt. Wellington Bett's Vale 147·15 42-55 Hickman, J.L. 1 1 04-Mar-54 Eagle Hawk Neck 147·55 43-00 Hickman, J.L. 2 1 16-Apr-54 Goulds Country near Lottah 148-05 41-15 Hickman, J.L. 6 2 17-Apr-S4 St Columba Falls 147-55 41-20 Hickman, J.L. 3 13-May-54 Eagle Hawk Neck 147·55 43-00 Hickman, J.L.

4 4 17-May-54 Burnie Fern Glade, nr 41-05 145-55 Hickman, J.L. 8 2 22-May-54 Tarraleah 42-20 146-25 Hickman, J.L. 2 24-May-54 Bronte Lyell Highway, 5 miles towards Hobart 42-15 146-35 Hickman, J.L. 1 1 24-MaY-54 Bronte , 5 mls towards Hobart 42-15 146-35 Hickman, J.L. 2 2 24-MaY-54 Lyell Highway Dee Bridge 42-15 146-35 Hickman, J.L. 8 2 26-May-54 Bronte Marlborough Highway 42-10 146-30 Hickman, J.L. 16 2 26-May-54 41-55 146-45 Hickman, J.L. 39 4 27-May-54 Tarraleah over pipeline 42-20 146-25 Hickman, J.L. 19 1 27-May-54 Tarraleah Butlers Gorge Road 42-20 146-25 Hickman, J.L. 13 4 28-MaY-54 Hellyer Gorge 41-20 145-35 Hickman, J.L. 5 2 14-Aug-54 East Risdon 42-50 147-20 Hickman, J.L. 8 1 14-Aug-54 Eagle Hawk Neck 43-00 147-55 Hickman, J.L. 4 2 14-Aug-54 Hobart Domain 42-50 147-20 Hickman, J.L. 5 1 17-Aug-54 Hobart Water-works Road 42-50 147-20 Hickman, J.L. 31 2 18-Aug-54 Parattah 42-20 147-25 Hickman, J.L. 4 2 18-Aug-54 Tunnack 42-25 147-30 Hickman, J.L. 11 4 24-Aug-54 Table Cape 41-00 145-45 Hickman, J.L. 9 5 24-Aug-54 Burnie, N.W. Tas. Fern Glade, Emu Road 41-05 145-55 Hickman, J.L. 4 2 25-Aug-54 Parrawe 41-20 145-35 Hickman, J.L. 9 1 13-0ct-54 Penguin lroncliff Road, Fern Dene 41-05 146-05 ? 13 1 21-Jul-55 Florentine Valley 42-35 146-25 Gilbert, J.M. 3 2 27-0ct-55 Mt Wellington 42-55 147-15 Hickman, J.L. 11 3 08-Nov-55 Collinsvale 42.50 147-05 Hickman, J.L. 9 1 11-Nov-55 Arve Valley 43-05 146-50 Hickman, J.L. 4 1 13-Nov-55 Mt Wellington 42-55 147-15 Hickman, J.L. 9 1 Apr. 1956 Eagle Hawk Neck 43-00 147-55 Hickman, J.L. 35 3 24-Jun-57 Lenah Valley New Town Falls 42-50 147-20 Hickman, J.L. 1 1 04-Aug-57 Tinderbox Tinderbox 43-05 147-20 Hickman, J.L. 2 1 24-Sep-58 ANM Road Bridge 42-50 146-35 Francombe, D.

5 2 1 Dec 1967 Port Davey Melaleuca Inlet 43-20 145-55 Dartnall, A.J. 25-Feb-71 Maria Creek 42.55 146-10 Tyler, P.A. 2 1 13-Mar-71 Mt Arthur 41-15 147-20 Dartnall, A.J. 4 1 10-Apr-71 Summit of Mt Mangana 43-25 147-15 Dartnall, A.J. 23 4 19-Aug-71 Mt. Wellington Shoo bridge Bend Track 42-55 147-15 Jamieson, B., & Bradbury, E. 1 1 19-5ep-71 Derby nr Old Breiseis Mine dam 41-10 147-50 Kershaw, R.C. 5 3 15-0ct-71 Mt Arthur 41-15 147-20 Dartnall, A.J. & Kershaw, R.C. 5 1 ? "Hobart and Straham" 42-50 147-20 ?

398 109 number of sites: 65 ( Mt Wellington: 10, Hobart: 7, Tarraleah: 5 )

6 TABLE 2 Collections made by Professor Jamieson in 1972 and donated to QVM in 1991 n spp date locality locality detail lat. & long.

9-Aug-72 Launceston, N. TAS 9 miles from Launceston on road to Scottsdale 41 27 S 147 10 E 14 2 9-Aug-72 Launceston, N. TAS Mt Barrow near ruined stone hut 41 27 S 147 10 E 16 2 9-Aug-72 Launceston, N. TAS Mt Barrow 41 27 S 147 10 E 55 2 9-Aug-72 Weldborough, N.E. TAS Weldborough Pass 41 12 S 147 54 E 34 1 10-Aug-72 St Marys, E. TAS Top of St Mary's Pass 41 35 S 148 12 E 14 1 10-Aug-72 St Helens, E. TAS St Helens to Hobart Road, 2 mls from St Helens 41 20 S 148 14 E 5 1 10-Aug-72 Hobart, S.E. TAS Foot of Mt Wellington 42 53 S 147 19 E 2 1 10-Aug-72 Hamilton, S.E. TAS Miena Road 0 10-Aug-72 Hamilton, S.E. TAS 8 miles from Hamilton along road to Bothwell 42 33 S 146 50 E 0 10-Aug-72 Bothwell, S.E.TAS 12 miles from Bothwell along road to Great Lake 6 1 10-Aug-72 Liapootah, C.P. TAS 1 mile east of Liapootah 0 S 0 E 0 11-Aug-72 Lake St Clair, W. T AS Banks of Lake St Clair 42 5S 146 5E 14 2 11-Aug-72 Queenstown, W. TAS Lyell Highway, 4 miles West of Mt Arrowsmith 42 5S 145 33 E 24 2 11-Aug-72 Queenstown, W. TAS Mt Arrowsmith 42 5S 145 33 E 4 2 12-Aug-72 Queenstown, W. TAS Nelson Valley Ck on Lyell Hwy, 14 mls E of Queenstown 42 5S 145 33 E 9 2 12-Aug-72 Queenstown, W. TAS 17 miles from Strahan along road to Queenstown 42 5S 145 33 E 10 1 13-Aug-72 Queenstown, W. TAS 7 miles from Queenstown along road to Burnie 42 5S 145 33 E 20 1 13-Aug-72 Zeehan, W. TAS , 4 miles N of Zeehan turnoff 41 54 S 145 21 E 19 1 13-Aug-72 Rosebery, W. TAS Murchison Highway, 4 miles West of Rosebery 41 47 S 145 34 E 2 1 13-Aug-72 Rosebery, W. TAS Mt Block on Murchison Highway 41 47 S 145 34 E 36 5 13-Aug-72 Tullah, N.W. TAS Murchison Highway, 15 miles N. of Tullah 41 44 S 145 37 E 111 7 13-Aug-72 Waratah, N.W. TAS Murchison Highway near Wandie River 41 27 S 145 32 E 4 1 13-Aug-72 Somerset, N.W. TAS 1 mile south of Somerset 41 3S 145 50 E

399 36 number of sites: 23 number of species: 19

7 was chosen on the basis that it supported a range of vegetation communities largely determined by altitude and aspect. Thirteen diverse sites were selected for study.

Two additional and significant opportunities to collect earthworms in remote locations arose in 1992 through the WEBS initiative of the Tasmanian Parks and Wildlife Service. This programme allowed myself and an assistant to spend a week based at each of Pelion Valley in February and at Melaleuca in March of that year. Earthworms were sampled at 14 localities within 10 km of the Pelion Rangers' Hut and at 10 localities within a 6 km radius of the airstrip at Melaleuca.

These and other directed and opportunistic collecting efforts during the two years preceding the National Estate-funded project allowed the accumulation of a further 950 specimens from 64 sites as summarised in Table 3.

In addition to these substantive collections, additional specimens arising from public enquiries and local surveys over the past four decades are shelved within the general collections of both the Queen Victoria Museum and the Tasmanian Museum. These specimens tend to be poorly preserved and poorly referenced, compared with the highly targetted collections assembled by the present author since 1990.

What was the aim of the project?

As stated in the grant application, the aim of the project was:

"To increase the level of knowledge of Tasmanian native earthworms from the present low level to a point at which it will be possible to evaluate the National Estate significance of the fauna and to assess the conservation status of each species. A guide book of the distribution and preference of each species will, for the first time, enable prediction of the impact of developments upon native earthworm species"

An additional implied aim was to attempt to reach a point at which it would be possible to conduct studies of the impact of forestry practices on earthworm populations and to assess their value as 'bio-indicators' of forest ecosystem health.

8 TABLE 3 Collections made by the author and associates 1990 - 1992

n spp date locality locality detail grid ref. alt. collectors 41 9 23 -Aug- 90 Weldborough, N.E. TAS Weld borough EQ 744 351 680 Mesibov, R. 79 6 29 -Aug- 90 Weldborough, N.E. TAS Weldborough EQ 744 353 650 Mesibov, R. 38 6 7 -Sep- 90 Weldborough, N.E. TAS Weldborough EQ 745 353 660 Mesibov, R. 55 4 8 -Sep- 90 Waratah, N.W. TAS Waratah CQ 704 066 690 Mesibov, R. 41 7 19 -Sep- 90 Waratah, N.W. TAS Waratah CQ 702 064 680 Mesibov, R. 42 5 23 -Sep- 90 Waratah, N.W. TAS Wa rata h CQ 703 065 670 Mesibov, R. 4 1 -Sep- 90 Weldborough, N.E. TAS EQ 756 395 Kingston, T.J. &Mesibov, R. 5 1 28 -Nov- 90 Mt. Arthur, N. TAS Patersonia Rivulet EQ 252 313 610 Mesibov, R. 24 4 6 -Dec- 90 Christmas Hills, N.W. TAS Christmas Hills CQ 309 667 60 Mesibov, R. 10 4 7 -Dec- 90 Christmas Hills, N.W. TAS CQ 341 702 20 Mesibov, R. 5 10 -Dec- 90 Roger River, N.W. TAS CQ 259 487 30 Mesibov, R. 14 3 16 -Dec- 90 Smithton, N.W. TAS Big Eel Ck CQ 180 303 190 Mesibov, R. 10 2 17 -Dec- 90 Trowutta Plateau, N.W. TAS CQ 330 483 210 Mesibov, R. 12 2 18 -Dec- 90 Frankland River Bridge, N.W. TAS Frankland River track CQ 251 313 110 Mesibov, R. 1 1 20 -Dec- 90 Balfour, N.W. T AS Frankland Plain CQ 239 402 160 Mesibov, R. 12 2 28 -Dec- 90 Roger River West N.W. TAS Montagu Swamp CQ 287 547 30 Mesibov, R. 11 2 29 -Dec- 90 Mt. Hazelton, N.W. TAS Mt Hazelton CQ 266 189 650 Mesibov, R. 4 1 29 -Dec- 90 Mt. Frankland, N.W. TAS Mt Frankland CQ 269 262 310 Mesibov, R. 6 1 15 -Apr- 91 Triabunna, E. TAS Buckland Military Area EP 688015 410 Mesibov, R. 3 1 16 -Apr- 91 Triabunna, E. TAS Mitchelmore's Ck EP 727 106 120 Mesibov, R. 3 1 17 -Apr- 91 Triabunna, E. TAS Buckland Military Area EN 696 961 150 Mesibov, R. 8 14 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 1 EQ 253 320 650 Mesibov, R. 4 1 14 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 2 EQ 255 327 550 Mesibov, R. 9 3 17 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 3 EQ 255 317 630 Mesibov, R. 9 1 17 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 4 EQ 251 308 660 Mesibov, R. 8 19 -May- 91 Mt. Arthur, N. TAS Mt. Arthu r site 5 EQ 224 325 520 Mesibov, R.

9 7 1 22 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 6 EO 234 305 1080 Mesibov, R. 19 3 22 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 7 EO 231 303 990 Mesibov, R. 2 1 18 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 8 EO 223 292 750 Mesibov, R. 32 4 18 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 9 EO 246 310 750 Mesibov, R. 50 3 21 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 10 EO 239 281 840 Mesibov, R. 6 3 18 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 11 EO 213 274 520 Mesibov, R. 1 1 21 -May- 91 Mt. Arthur, N. TAS Mt. Arthur site 12 EO 209 282 480 Mesibov, R. 2 1 11 -Aug- 91 Sideling Range, N.E. TAS Kingston, T.J. 1 1 14 -Aug- 91 Cape Naturalist, N.E. TAS Site 3 Kingston, T.J. & McGowan, L.F. 9 1 14 -Aug- 91 Cape Naturalist, N.E. TAS Top Camp Kingston, T.J. &McGowan, L.F. 4 1 15 -Aug- 91 Waterhouse, N.E. TAS Little Waterhouse EO 516 751 Kingston, T.J. 5 3 27 -Aug- 91 Gog Range, N. TAS Gog Range site 2 DO 534056 Kingston, T.J. 4 1 27 -Aug- 91 Gog Range, N. TAS Gog Range site 1 DO 543058 Kingston, T.J. 2 1 13 -Dec- 91 Gog Range, N. TAS Gog Range DO 548059 Mesibov, R. 4 1 14 -Dec- 91 Mt. Barrow, N.E. TAS Mt. Barrow EO 348 783 1295 Mesibov, R. 5 1 16 -Dec- 91 Gog Range, N. TAS Gog Range DO 461058 530 Mesibov, R. 3 1 16 -Jan- 92 Pioneer, N.E. TAS Pioneer EO 782 512 Kingston, T.J. & Laffan, M. 27 4 10 -Feb- 92 Pelion Valley, W. TAS Mt. Oakleigh DP 208 705 985 Kingston, T.J. &M.E. 13 4 10 -Feb- 92 Pelion Valley, W. TAS Mt. Oakleigh DP 209 700 870 Kingston, T.J. &M.E. 20 6 10 -Feb- 92 Pelion Valley, W. TAS Mt. Oakleigh DP 209 702 895 Kingston, T.J. &M.E. 3 2 11 -Feb- 92 Pelion Valley, W. TAS track DP 211 655 980 Kingston, T.J. &M.E. 12 3 11 -Feb- 92 Pelion Valley, W. TAS Pelion Gap track DP 213 653 1010 Kingston, T.J. & M.E. 9 3 11 -Feb- 92 Pelion Valley, W. TAS Pelion Gap track DP 217 649 1120 Kingston, T.J. &M.E. 2 1 12 -Feb- 92 Pelion Valley, W. TAS buttongrass plain DP 201 687 845 Kingston, T.J. &M.E. 15 5 12 -Feb- 92 Pelion Valley, W. TAS nr Old Pelion Hut DP 204 684 860 Kingston, T.J. &M.E. 11 5 12 -Feb- 92 Pelion Valley, W. TAS DP 209 660 940 Kingston, T.J. &M.E. 23 5 13 -Feb- 92 Pelion Valley, W. TAS , N.W. side DP 173 674 725 Kingston, T.J. &M.E. 19 7 13 -Feb- 92 Pelion Valley, W. TAS Forth River, S.E. side DP 177 675 730 Kingston, T.J. &M.E. 3 1 13 -Feb- 92 Pelion Valley, W. TAS Forth River, S.E. side DP 180 678 760 Kingston, T.J. & M.E. 16 3 13 -Feb- 92 Pelion Valley, W. TAS Forth River, S.E. side DP 196 683 875 Kingston, T.J. &M.E.

10 19 5 14 -Feb- 92 Pelion Valley, W. TAS Ann River track DP 194 710 750 Kingston, T.J. & ME 10 2 1 -Mar- 92 Melaleuca area, S.W. TAS nr Half-woody Hill DM 339 909 15 Kingston, T.J. & McGowan, LF. 9 1 3 -Mar- 92 Melaleuca area, S.W. TAS Denny King's garden DM 319 922 2 Kingston, T.J. & McGowan, LF. 23 2 4 -Mar- 92 Melaleuca area, S.W. TAS Celery Top Island DM 309 977 8 Kingston, T.J. & McGowan, LF. 11 2 4 -Mar- 92 Melaleuca area, S.W. TAS edge of Melaleuca Lagoon DM 321 921 1 Kingston, T.J. & McGowan, LF. 28 3 5 -Mar- 92 Melaleuca area, S.W. TAS DM 328 904 10 Kingston, T.J. & McGowan, LF. 61 6 5 -Mar- 92 Melaleuca area, S.W. TAS Half-woody Hill DM 338 889 80 Kingston, T.J. & McGowan, LF. 3 1 23 -Apr- 92 Scottsdale, N.E. TAS Scottsdale EQ 434 432 Kingston, T.J.

11 TABLE 4 Collections made under the present study n spp date locality locality detail grid ref. alt. collectors

13 2 2·May- 92 Retreat, N. TAS EO 144 481 135 Kingston, T.J. & D'orazio, R. 22 2 2·May- 92 Retreat, N. TAS EO 156 473 130 Kingston, T.J. & D'orazio, R. 40 3 2·May- 92 Retreat, N. TAS EO 160 451 160 Kingston, T.J. & D'orazio, R. 24 4 3 -Jun- 92 Springfield South, N.E. TAS off East Diddleum Road EO 388 287 560 Kingston, T.J. & D'orazio, R. 15 2 9 -Jun- 92 Golconda, N. TAS EO 264 438 70 Kingston, T.J. & D'orazio, R. 34 3 9 -Jun- 92 Springfield, N.E. TAS off Upper Brid Road EO 380 307 330 Kingston, T.J. & D'orazio, R. 5 3 9 -Jun- 92 Jetsonville, N.E. TAS EO 393 518 70 Kingston, T.J. & D'orazio, R. 8 1 15 -Jun- 92 Nabowla, N. TAS EO 263 383 260 Kingston, T.J. & D'orazio, R. 86 8 15 -Jun- 92 Sideling Range, N.E. TAS EO 344 353 530 Kingston, T.J. & D'orazio, R. 14 4 16 -Jun- 92 Mt. Barrow, N.E. TAS Mt. Barrow site 1 EO 317 223 500 Kingston, T.J. & D'orazio, R. 8 2 16 -Jun- 92 Mt. Barrow, N.E. TAS Mt. Barrow site 2 EO 353 212 750 Kingston, T.J. & D'orazio, R. 3 2 16 -Jun- 92 Mt. Barrow, N.E. TAS Mt. Barrow site 3 EO 354 193 1240 Kingston, T.J. & D'orazio, R. 26 6 16 -Jun- 92 Mt. Barrow, N.E. TAS Mt. Barrow site 4 EO 358 195 1020 Kingston, T.J. & D'orazio, R. 8 1 16 -Jun- 92 Mt. Barrow, N.E. TAS Mt. Barrow site 5 EO 358 202 895 Kingston, T.J. & D'orazio, R. 10 2 17 -Jun- 92 Merthyr Park, N. TAS EO 160 342 180 Kingston, T.J. & D'orazio, R. 15 2 17 -Jun- 92 Hollybank, N. TAS EO 179 270 300 Kingston, T.J. & D'orazio, R. 17 2 17 -Jun- 92 Lilydale Falls, N.E. TAS EO 179 355 180 Kingston, T.J. & D'orazio, R. 6 2 17 -Jun- 92 Prossers Forest, N. T AS EO 194 205 420 Kingston, T.J. & D'orazio, R. 23 5 22 -Jun- 92 Mt. Victoria, N.E. TAS Mt. Victoria F.R. site 1 EO 662 225 720 Kingston, T.J. & D'orazio, R. 51 3 22 -Jun- 92 Mt. Victoria, N.E. TAS Mt. Victoria F.R. site 2 EO 691 226 790 Kingston, T.J. & D'orazio, R. 25 5 22 -Jun- 92 Mt. Victoria, N.E. TAS Mt. Victoria F. R. site 3 EO 718 225 810 Kingston, T.J. & D'orazio, R. 30 8 23 -Jun- 92 Mathinna, E. TAS Griffin F.R. EO 687 088 Kingston, T.J. & D'orazio, R. 73 9 23 -Jun- 92 Mathinna, E. TAS Evercreech F.R. EO 812 162 350 Kingston, T.J. & D'orazio, R. 26 5 24 -Jun- 92 Mt. Maurice, N.E. TAS Mt. Maurice site 1 EO 503 263 940 D'orazio, R. & McGowan, L.F. 9 3 24 -Jun- 92 Mt. Maurice, N.E. TAS Mt. Maurice site 2 EO 505 262 910 D'orazio, R. & McGowan, L.F. 28 6 24 -Jun- 92 Mathinna Falls, N.E. TAS Mathinna Falls Reserve EO 748 162 470 Kingston, T.J. & D'orazio, R. 21 3 25 -Jun- 92 Tombstone Creek F.R., N.E. TAS EO 577 163 640 D'orazio, R. & Cooper, M. 24 5 29 -Jun- 92 Dazzler Range, N. TAS Kerrisons Road site 1 DO 755 376 415 D'orazio, R. & Mitchell, A.

12 30 6 29 -Jun- 92 Dazzler Range, N. TAS Kerrisons Road site 2 DO 756 389 510 D'orazio, R. & Mitchell, A. 33 4 29 -Jun- 92 Dazzler Range, N. TAS Horders Rd DO 764 349 330 D'orazio, R. & Mitchell, A. 13 4 29 -Jun- 92 Dazzler Range, N. TAS Tattersalls Road DO 773 359 440 D'orazio, R. & Mitchell, A. 6 1 29 -Jun- 92 Dazzler Range, N. TAS site 5 EO 753 368 340 D'orazio, R. & Mitchell, A. 35 4 30 -Jun- 92 Dalgarth F.R., N. TAS site 1 DO 707 343 120 Kingston, T.J. & D'orazio, R. 45 3 30 -Jun- 92 Dalgarth F.R., N. TAS site 2 DO 722 325 0 Kingston, T.J. & D'orazio, R. 59 5 30 -Jun- 92 Dalgarth F.R., N. TAS Wallaby Ck DO 734 329 0 Kingston, T.J. & D'orazio, R. 26 6 2 -Jul- 92 Fairy Glade S.R .. N. TAS DP 766 883 740 D'orazio, R. & McGowan, L.F. 35 7 2 -Jul- 92 Liffey Falls, N. TAS DP 763 827 920 D'orazio, R. & McGowan, L.F. 32 4 2 -Jul- 92 Liffey Falls, N. TAS DP 813 845 520 D'orazio, R. & McGowan, L.F. 0 0 6 -Jul- 92 Cluan Tier, N. TAS Wand ilia Road DP 805 946 425 D'orazio, R. & Mitchell, A. 5 2 6 -Jul- 92 Cluan Tier, N. TAS Myrtle Ck Road DP 872 878 520 D'orazio, R. & Mitchell, A. 23 5 7 -Jul- 92 Legerwood, N.E. TAS Cuckoo Falls EO 516 343 415 D'orazio, R. 30 5 7 -Jul- 92 Legerwood, N.E. TAS Hogarth Road EO 521 356 290 D'orazio, R. 17 1 7 -Jul- 92 Legerwood, N.E. TAS Cuckoo Hill Road EO 543 338 455 D'orazio, R. 14 2 9 -Jul- 92 Liffey, N. TAS Coalmine Ck DP 833 848 510 D'orazio, R. 9 1 9 -Jul- 92 Drys Bluff, N. TAS DP 867 837 450 D'orazio, R. 32 3 16 -Jul- 92 Blackwood Creek, N. TAS Westons Rivulet DP 895 758 375 D'orazio, R. & Cooper, M. 0 0 16 -Jul- 92 Caseyville, N. TAS Abraham Ck EP 38 636 360 D'orazio, R. & Cooper, M. 22 3 20 -Jul- 92 Pioneer, N.E. TAS Royston Ck EO 828 493 215 D'orazio, R. & Cooper, M. 17 2 20 -Jul- 92 Pioneer, N.E. TAS Old Chum Dam EO 883 539 110 D'orazio, R. & Cooper, M. 63 6 20 -Jul- 92 Mt. Cameron, N.E. TAS Water Race Reserve EO 886 483 175 D'orazio, R. & Cooper, M. 23 3 21 -Jul- 92 Ringarooma, N.E. TAS Mt. Paris Road EO 677 376 540 D'orazio, R. & Cooper, M. 29 4 21 -Jul- 92 Moorina, N.E. TAS Frome Road EO 762 443 420 D'orazio, R. & Cooper, M. 31 3 21 -Jul- 92 Weldborough Pass, N.E. TAS Myrtle F.R. EO 777 372 440 D'orazio, R. & Cooper, M. 72 6 21 -Jul- 92 Weldborough, N.E. TAS Emu Road EO 781 408 540 D'orazio, R. & Cooper, M. 0 0 23 -Jul- 92 Launceston, N. TAS Punchbowl Recreation Reserve EO 139 103 30 D'orazio, R. & Cooper, M. 6 1 23 -Jul- 92 Launceston, N. TAS Carr Villa Floral Reserve EO 144 092 D'orazio, R. & Cooper, M. 10 1 27 -Jul- 92 Scamander, E. TAS Scamander F.R. FO 015 123 100 D'orazio, R. & Cooper, M.

13 21 5 27 -Jul- 92 Toms Gully, E. TAS FO 900 218 320 D'orazio, R. & Cooper, M. 19 3 28 -Jul- 92 Peters Road, E. T AS EO 939 445 170 D'orazio, R. & Cooper. M. 26 4 28 -Jul- 92 Wild Pig Hill, E. TAS EO 962 489 160 D'orazio, R. & Cooper, M. 11 2 28 -Jul- 92 Badger Marsh, N.E. TAS EO 975 368 120 D'orazio, R. & Cooper, M. 33 4 29 -Jul- 92 Mt. Nicholas Road, E.TAS EP 900 995 520 D'orazio, R. & Cooper. M. 41 5 29 -Jul- 92 Pyengana, N.E. TAS Intake Bridge EO 781 260 360 D'orazio, R. & Cooper, M. 20 3 29 -Jul- 92 Trafalgar Flat, E. TAS EO 875 178 300 D'orazio, R. & Cooper. M. 7 1 3-Aug- 92 Fingal, E. TAS Valley Road EP 875 895 500 D'orazio, R. & Cooper, M. 1 1 3-Aug- 92 Mt. Puzzler F.R., E. TAS Meadstone Falls Track EP 892 791 480 D'orazio, R. & Cooper, M. 17 3 3·Aug- 92 Fingal, E. TAS Valley Road EP 894 836 760 D'orazio, R. & Cooper, M. 3 2 3-Aug- 92 Fingal. E. TAS Valley Road EP 985 875 500 D'orazio, R. & Cooper, M. 24 2 4-Aug- 92 St Marys Pass S.R., E. TAS Lower GermanTown Road FP 002 989 220 D'orazio, R. & Cooper, M. 19 2 4-Aug- 92 lrishtown, N.W. TAS St Pat ricks Head S.R. FP 026 988 430 D'orazio, R. & Cooper, M. 4 2 5-Aug- 92 Hardings Falls F.R., E. TAS EP 914 663 260 D'orazio, R. & Cooper. M. 35 4 5-Aug- 92 Apsley Myrtle F.R.. E. TAS EP 932 719 450 D'orazio, R. & Cooper, M. 5 5 ·Aug- 92 Bicheno, E. TAS Swilly Marsh EP 956 623 280 D'orazio, R. & Cooper, M. 21 2 5-Aug- 92 Chain of lagoons, E. T AS 'E' Road, Piccaninny Ck bridge FP 029 847 310 D'orazio, R. & Cooper, M. 25 3 10 -Aug- 92 Royal George, E. T AS Meetus Falls EP 731 552 600 D'orazio, R. & Cooper, M. 23 4 10-Aug- 92 Lake Leake, E. TAS Lost Falls F.R. EP 746 447 470 D'orazio, R. & Cooper, M. 16 2 10-Aug- 92 Royal George, E. T AS West Swan River EP 790 605 540 D'orazio, R. & Cooper, M. 38 5 11 -Aug- 92 Tooms Lakes, E. TAS Tooms White Gum F.R. EP 701 226 600 D'orazio, R. & Cooper, M. 25 5 11 -Aug- 92 Brookerana FR., E. TAS EP 709 193 590 D'orazio, R. & Cooper, M. 55 6 11 -Aug- 92 Tom Legges Tier, E. TAS McKays Road EP 728 302 600 D'orazio, R. & Cooper, M. 28 4 17 -Aug- 92 Colebrook, S.E. TAS Coal River Gorge Nature Reserve EN 325 965 310 D'orazio, R. & Cooper, M. 16 2 17 -Aug- 92 Colebrook, S.E. TAS Spinning Gum F.R. EN 393 950 460 D'orazio, R. & Cooper, M. 20 3 18-Aug- 92 Dysart. Midlands, T AS Barren Rock Falls F.R. EN 208 867 350 D'orazio, R. & Cooper, M. 0 0 18-Aug- 92 Bagdad, Midlands, TAS Chauncy Vale Wildlife Sanctuary EN 237 820 330 D'orazio, R. & Cooper. M. 3 19-Aug- 92 Pipers Brook, N. TAS Pipers Brook Road EO 152 537 100 D'orazio, R. & Cooper, M. 0 0 19-Aug- 92 Waverley, Launceston, N. TAS Distillary Ck Reserve EO 153 132 60 D'orazio, R. & Cooper, M.

14 17 1 19-Aug- 92 Pipers Brook, N. T AS State Forest. Bridport Highway EO 175 553 80 D'orazio, R. & Cooper, M. 25 3 24 -Aug- 92 Frankford, N. TAS Mt. Careless State Forest DO 802 283 420 D'orazio, R. & Cooper, M. 9 2 24 -Aug- 92 Holwell, N. TAS Holwell Gorge DO 804 304 300 D'orazio, R. & Cooper, M. 13 2 24-Aug- 92 FourSprings,N.TAS Four Springs Forest Park DO 863 199 180 D'orazio, R. & Cooper, M. 17 3 24-Aug- 92 Notley Gorge, N. TAS DO 926 215 240 D'orazio, R. & Cooper, M. 26 4 25-Aug- 92 Railton, N.W. TAS RedwaterCk DO 487 195 200 D'orazio, R. & Cooper, M. 21 3 25 -Aug- 92 Railton, N.W. TAS Henry Somerset ConselVation Area DO 508 305 45 D'orazio, R. & Cooper, M. 44 4 25-Aug- 92 Kimberley, N.W. TAS Richland Hill DO 512 168 180 D'orazio, R. & Cooper, M. 34 4 26-Aug- 92 Forth, N.W. TAS Champion Nature ReselVe DO 372 372 40 D'orazio, R. & Cooper, M. 6 2 26-Aug- 92 Eugenana, N.W. TAS Arboretum Nature Trail DO 414 357 50 D'orazio, R. & Cooper, M. 37 5 26-Aug- 92 Paradise, N.W. TAS MinnowCk DO 447 086 260 D'orazio, R. & Cooper, M. 12 4 31 -Aug- 92 Forth Falls S.R., N.W. TAS Lake Barrington Rd. DO 333 185 220 D'orazio. R. & Cooper. M. 15 4 31-Aug- 92 Lower Wilmot. N.W. TAS Ingram Ck, Wilmot Road DO 346 277 240 D'orazio. R. & Cooper, M. 29 2 31-Aug- 92 Devonport, N.W. TAS Bella - Macargee Falls DO 402 376 100 D'orazio, R. & Cooper, M. 10 1 1 -Sep- 92 , N. TAS Baldocks Cave State Forest DP 444 958 430 D'orazio, R. 7 2 1 -Sep- 92 Needles, N. TAS Lobster Falls track DO 598 008 360 D'orazio, R. 11 2 2-Sep- 92 Mole Creek, N. TAS Croesus Cave S.R. DP 352 966 300 D'orazio, R. 0 0 2-Sep- 92 Mole Creek, N. TAS King Solomons Cave S.R. DP 371 997 440 D'orazio, R. 8 2 2-Sep- 92 Mole Creek, N. TAS Sensation Gorge Scenery ReselVe DP 442 992 290 D'orazio, R. 70 6 7 -Sep- 92 Tasman Peninsula, S.E. T AS Pirates Road, Taranna EN 711 476 70 Kingston, T.J. & D'orazio, R. 32 3 7 -Sep- 92 Tasman Peninsula, S.E. TAS Pirates Road, Taranna EN 723 338 105 Kingston, T.J. & D'orazio, R. 38 5 7 -Sep- 92 Tasman Peninsula, S.E. TAS Hylands Road EN 752 450 120 Kingston, T.J. & D'orazio, R. 28 4 7 -Sep- 92 Tasman Peninsula, S.E. TAS Lizard Hill EN 756 418 230 Kingston, T.J. & D'orazio, R. 25 4 8 -Sep- 92 Tasman Peninsula, S.E. TAS Mt Arthur EN 654 204 375 Kingston, T.J. & D'orazio, R. 28 5 8 -Sep- 92 Tasman Peninsula, S.E. TAS Long Bay Ck, Fortescue Road EN 708 269 20 Kingston, T.J. & D'orazio, R. 26 4 8 -Sep- 92 Tasman Peninsula, S.E. TAS Baits Road EN 723 301 140 Kingston, T.J. & D'orazio, R. 40 2 8 -Sep- 92 Tasman Peninsula, S.E. TAS Cape Pillar Track, Fortescue Road EN 761 228 70 Kingston, T.J. & D'orazio, R. 33 4 9-Sep- 92 Tasman Peninsula, S.E. TAS Badger Ck EN 587 310 190 Kingston, T.J. & D'orazio, R. 53 5 9-Sep- 92 Tasman Peninsula, S.E. TAS Benjafields Ridge EN 607 228 260 Kingston, T.J. & D'orazio, R.

15 39 5 9-Sep- 92 Tasman Peninsula, S.E. TAS Koonya State Forest EN 647 269 315 Kingston, T.J. & D'orazio, R. 9 2 15-Sep- 92 Mole Creek, N. TAS Marakoopa Cave S.R. DP 405 967 500 D'orazio, R. 28 3 15-Sep- 92 Chudleigh, N. TAS Westmorland Falls Track DP 493 922 460 D'orazio, R. 5 3 16-Sep- 92 Mole Creek, N. TAS Martha Ck, Mersey Forest Road DP 352 922 400 D'orazio, R. 40 5 16-Sep- 92 Mole Creek, N. TAS Urks Loop, Mersey Forest Road DP 367 975 560 D'orazio, R. 6 2 16 ·Sep- 92 Mole Creek, N. T AS King Solomons Cave S.R. DP 373 993 420 D'orazio, R. 25 4 28-Sep- 92 Bruny Island, North, S.E. TAS Gravel Reserve No. 0133 EN 290 287 60 D'orazio, R. & Cooper, M. 24 4 28-Sep- 92 Bruny Island, North, S.E. TAS McCrackens Ck, Missionary Road EN 294 244 20 D'orazio, R. & Cooper, M. 1 1 28 ·Sep- 92 Bruny Island, North, S.E. TAS Bruny Island Neck Game Reserve EN 306 134 10 D'orazio, R. & Cooper, M. 12 2 29 ·Sep- 92 Bruny Island, South, S.E. TAS Goulds Ck, Conley Road EM 198 997 150 D'orazio, R. & Cooper, M. 13 2 29-Sep- 92 Bruny Island, South, S.E. TAS Clennetts Road EM 219 931 75 D'orazio, R. & Cooper, M. 24 3 29 ·Sep- 92 Bruny Island, South, S.E. TAS Saintys Ck, Coolangatta Road EM 222 981 365 D'orazio, R. & Cooper, M. 18 2 29 ·Sep- 92 Bruny Island, South, S.E. TAS Staffords Road EM 233 905 145 D'orazio, R. & Cooper, M. 26 3 29-Sep- 92 Bruny Island, South, S.E. TAS South Bruny Range EM 251 909 350 D'orazio, R. & Cooper, M. 39 2 29 -Sep- 92 Bruny Island, South, S.E. TAS Captain Cook Ck EM 258 952 175 D'orazio, R. & Cooper, M. 37 4 30 -Sep- 92 Bruny Island, South, S.E. TAS La Billiardiere S.R. EM 142 866 90 D'orazio, R. & Cooper, M. 40 4 30-Sep- 92 Bruny Island, South, S.E. TAS Resolution Road EN 241 012 125 D'orazio, R. & Cooper, M. 5 1 5 -Oct- 92 Mole Creek, N. TAS Arm River F.R. DP 332 839 460 D'orazio, R. & Cooper, M. 29 4 5 -Oct- 92 Mole Creek, N. TAS Lake MacKenzie Road site 1 DP 362 942 540 D'orazio, R. & Cooper, M. 15 4 5 -Oct- 92 Mole Creek, N. TAS Lake MacKenzie Road site 2 DP 373 924 700 D'orazio, R. & Cooper, M. 27 4 5 -Oct- 92 Mole Creek, N. TAS Snake Ck Road DP 391 895 590 D'orazio, R. & Cooper, M. 10 2 5 -Oct- 92 Mole Creek, N. TAS Devils Gullet S.R. DP 443 876 1140 D'orazio, R. & Cooper, M. 10 1 6 -Oct- 92 Lake Rowallen, N. TAS Mersey White Water F. R. DP 351 808 445 D'orazio, R. & Cooper, M. 3 1 6 -Oct- 92 Lake Rowallen, N. TAS Dublin Road DP 363 802 650 D'orazio, R. & Cooper, M. 10 1 6 -Oct- 92 Lake Rowallen, N. TAS Fish River Road DP 365 749 720 D'orazio, R. & Cooper, M. 11 4 6 -Oct- 92 Lake Rowallen, N. TAS Dublin Road site 1 DP 381 825 590 D'orazio, R. & Cooper, M. 6 1 6 -Oct- 92 Lake Rowallen, N. TAS Dublin Road site 2 DP 385 833 575 D'orazio, R. & Cooper, M. 26 3 6 -Oct- 92 Lake Rowallen, N. TAS Little Fisher River Road DP 419 824 650 D'orazio, R. & Cooper, M. 44 4 12 -Oct- 92 Geeveston, S.E. TAS Peppers Road ON 857 252 260 D'orazio, R. & Cooper, M.

16 34 3 12 -Oct- 92 Geeveston, S.E. TAS Lidgerwood Ck ON 871 272 190 D'orazio, R. & Cooper, M. 45 5 12 -Oct- 92 Geeveston, S.E. TAS Bermuda Road ON 913 329 520 D'orazio, R. & Cooper, M. 40 6 12 -Oct- 92 Geeveston, S.E. TAS Bermuda Hill ON 918 320 520 D'orazio, R. & Cooper, M. 30 5 13 -Oct- 92 Geeveston, S.E. T AS South Weld F.R.. ON 722 366 270 D'orazio, R. & Cooper, M. 48 7 13 -Oct- 92 Geeveston, S.E. TAS South Weld Road. ON 755 347 235 D'orazio, R. & Cooper, M. 33 4 13 -Oct- 92 Geeveston, S.E. TAS Picton River ON 762 213 90 D'orazio, R. & Cooper, M. 52 7 13 -Oct- 92 Geeveston, S.E. TAS Tahune F.R. ON 778 286 60 D'orazio, R. & Cooper, M. 47 3 13 -Oct- 92 Geeveston, S.E. TAS Edwards Road ON 834 262 150 D'orazio, R. & Cooper, M. 21 3 14 -Oct- 92 Geeveston, S.E. TAS Hartz Mountain National Park ON 809 171 700 D'orazio, R. & Cooper, M. 13 3 14 -Oct- 92 Geeveston, S.E. TAS Hartz Road ON 812 169 680 D'orazio, R. & Cooper, M. 17 2 14 -Oct- 92 Geeveston, S.E. TAS KeoghsCk ON 828 218 180 D'orazio, R. & Cooper, M. 21 3 14 -Oct- 92 Geeveston, S.E. TAS Harveys Ck ON 854 186 520 D'orazio, R. & Cooper, M. 14 3 19 -Oct- 92 Dover, S.E. TAS Esperance River Road ON 899 082 140 D'orazio, R. & Cooper, M. 20 4 19 -Oct- 92 Dover, S.E. TAS Hermons Road ON 902 128 550 D'orazio, R. & Cooper, M. 24 3 19 -Oct- 92 Dover, S.E. TAS Riawunna Road ON 906 162 225 D'orazio, R. & Cooper, M. 26 2 19 -Oct- 92 Dover, S.E. TAS Esperance F.R. ON 928 590 75 D'orazio, R. & Cooper, M. 27 3 20 -Oct- 92 Dover, S.E. TAS Hastings Caves S.R. OM 878 955 85 D'orazio, R. & Cooper, M. 11 2 20 -Oct- 92 Dover, S.E. TAS South Lune Road OM 888 897 80 D'orazio, R. & Cooper, M. 11 2 20 -Oct- 92 Dover, S.E. TAS Ckton Road OM 890 952 70 D'orazio, R. & Cooper, M. 34 4 20 -Oct- 92 Dover, S.E. TAS Ckton Rivulet OM 922 983 110 D'orazio, R. & Cooper, M. 18 4 20 -Oct- 92 Dover, S.E. TAS Ckton Road ON 938 008 155 D'orazio, R. & Cooper, M. 43 8 21 -Oct- 92 Dover, S.E. TAS Stubbs Link Road ON 962 077 155 D'orazio, R. & Cooper, M. 26 3 21 -Oct- 92 Dover, S.E. T AS Wobbly Ck bridge ON 963 116 200 D'orazio, R. & Cooper, M. 33 5 21 -Oct- 92 Dover, S.E. TAS Hopetoun F.R. ON 971 051 190 D'orazio, R. & Cooper, M. 25 2 31 -Oct- 92 Flinders Island, N.E. TAS West End Road, Leeka ER 677 837 Kingston, T.J. & M.E. 5 1 31 -Oct- 92 Flinders Island, N.E. TAS West End Road, Leeka ER 705 825 Kingston, T.J. & M.E. 3 1 1·Nov- 92 Flinders Island, N.E. TAS Big River Kingston, T.J. & M.E. 6 2 1 ·Nov- 92 Flinders Island, N.E. TAS Mt. Strzelecki Kingston, T.J. & M.E. 56 4 2·Nov- 92 Flinders Island, N.E. TAS Strzelecki National Park ER 908 480 Kingston, T.J. & M.E.

17 16 2 2·Nov- 92 Flinders Island, N.E. TAS Mt. Strzelecki ER 925 485 Kingston, T.J. &M.E. 4 1 2·Nov- 92 Flinders Island, N.E. TAS Mt. Strzelecki Kingston, T.J. & M.E. 30 3 3·Nov- 92 Flinders Island, N.E. TAS Mt. Strzelecki ER 899 488 Kingston, T.J. & M.E. 5 2 3·Nov- 92 Flinders Island, N.E. TAS Mt. Strzelecki Kingston, T.J. &M.E. 16 3 4·Nov- 92 Flinders Island, N.E. TAS Brougham Sugarloaf ER 666 845 Kingston, T.J. & M.E. 11 2 4·Nov- 92 Flinders Island, N.E. TAS Walkers Lookout ER 918 653 350 Kingston, T.J. &M.E. 31 3 4·Nov- 92 Flinders Island, N.E. TAS Walkers Lookout ER 921 653 410 Kingston, T.J. &M.E. 22 2 4·Nov- 92 Flinders Island, N.E. TAS Middle Patriarch FR 014 730 120 Kingston, T.J. &M.E. 12 2 5·Nov- 92 Flinders Island, N.E. TAS 5 mile Road ER 848 830 Kingston, T.J. &M.E. 35 4 16·Nov- 92 Judbury, S.E. TAS Denison Road DN 802 416 160 D'orazio, R. &Cooper, M. 26 5 16·Nov- 92 Judbury, S.E. TAS Link Road DN 820 427 140 D'orazio, R. & Cooper, M. 14 4 16·Nov- 92 Judbury, S.E. TAS Russell Road site 1 DN 779 501 360 D'orazio, R. & Cooper, M. 13 2 16·Nov- 92 Judbury, S.E. TAS Russell Road site 2 DN 820 479 400 D'orazio, R. &Cooper, M. 15 2 17 ·Nov- 92 Mt. Wellington, S.E. TAS Big Bend Trail EN 173 518 1090 D'orazio, R. & Cooper, M. 22 4 17 ·Nov- 92 Mt. Wellington, S.E. TAS Huon Rd EN 185 445 420 D'orazio, R. &Cooper, M. 18 6 17 ·Nov- 92 Mt. Wellington, S.E. TAS Pinnacle Rd EN 192 512 1030 D'orazio, R. & Cooper, M. 16 2 17 ·Nov- 92 Mt. Wellington, S.E. TAS Badfords Track EN 199 485 700 D'orazio, R. & Cooper, M. 12 3 18·Nov- 92 Glenorchy, S.E. TAS Humphreys Track EN 192 547 250 D'orazio, R. & Cooper, M. 15 3 18·Nov- 92 Glenorchy, S.E. TAS Lena Valley Road EN 198 532 390 D'orazio, R. & Cooper, M. 11 3 18·Nov- 92 Glenorchy, S.E. TAS Water Reserve &Wildlife Sanctuary EN 202 532 320 D'orazio, R. &Cooper, M. 3 2 18·Nov- 92 Glenorchy, S.E. TAS Tolosa Road, Tolosa Park EN 206 537 200 D'orazio, R. & Cooper, M. 24 3 23·Nov- 92 Mt. Roland, N.W. TAS Cethana Road DQ 284 087 360 D'orazio, R. &Gittus, M. 5 1 23 ·Nov- 92 Mt. Roland, N.W. TAS Track off O'Neils Road. DQ 329 072 420 D'orazio, R. &Gittus, M. 18 3 23·Nov- 92 Mt. Roland, N.W. TAS O'Neils Road DQ 353 081 410 D'orazio, R. &Gittus, M. 27 3 23 ·Nov- 92 Mt. Roland, N.W. TAS Claude Road DQ 381 126 235 D'orazio, R. &Gittus, M. 23 3 23·Nov- 92 Mt. Roland, N.W. TAS Currawong Road DQ 406 115 390 D'orazio, R. & Gittus, M. 32 6 24·Nov- 92 Mt. Roland, N.W. TAS Cockatoo Road DQ 304 051 640 D'orazio, R. &Gittus, M. 30 4 24 ·Nov- 92 Mt. Roland, N.W. TAS Olivers Road DQ 344 021 715 D'orazio, R. & Gittus, M. 10 3 24·Nov- 92 Mt. Roland, N.W. TAS Liena Road DQ 389 004 455 D'orazio, R. &Gittus, M.

18 39 4 24·Nov- 92 Mt. Roland, N.W. TAS Belstone Road DQ 429 078 300 D'orazio, R & Gittus, M. 38 4 24·Nov- 92 Mt. Roland, N.W. TAS Union Bridge Road DQ 446 088 240 D'orazio, R & Gittus, M. 34 4 25·Nov- 92 Nietta, N.W. TAS Jean Brook Reserve DQ 193 116 575 D'orazio, R & Gittus, M. 11 2 25·Nov- 92 Lake Barrington, N.W. TAS Billet Ck Nature Walk. DQ 325 133 370 D'orazio, R &Gittus, M. 16 2 25·Nov- 92 Lower Barrington, N.W. TAS Lake Paloona Rd DQ 373 298 75 D'orazio, R & Gittus, M. 1 1 19 -Apr- 93 Yolla, N.W. TAS Blackwell Road site 1 CQ 785 351 250 D'orazio, R & Soccol, D. 10 3 19 -Apr- 93 Yolla, N.W. TAS Blackwell Road site 2 CQ 790 352 250 D'orazio, R &Soccol, D. 3 19 -Apr- 93 Calder, N.W. TAS West Calder Road CQ 839 517 110 D'orazio, R &Soccol, D. 10 3 19 -Apr- 93 Calder, N.W. TAS Oldina F.R CQ 876 595 70 D'orazio, R & Soccol, D. 10 3 19 -Apr- 93 Calder, N.W. TAS Tram Road Picnic Area CQ 892 572 34 D'orazio, R &Soccol, D. 27 5 20 -Apr- 93 Trowutta, N.W. TAS Milkshake F.R CQ 462 483 160 D'orazio, R & Soccol, D. 18 4 20 -Apr- 93 Trowutta, N.W. TAS Tayatea Road CQ 481 535 95 D'orazio, R &Socco!, D. 27 3 20 .Apr- 93 Mawbanna, N.W. TAS Dip Road CQ 576 661 70 D'orazio, R & Socco!, D. 22 5 20 -Apr- 93 Lileah, N.W. TAS Daisies Road CQ 624 545 120 D'orazio, R &Soccol, D. 20 2 20 -Apr· 93 Mawbanna, N.W. TAS Newhaven Road site 1 CQ 655 603 150 D'orazio, R &Socco!, D. 6 1 20 ·Apr- 93 Mawbanna, N.W. TAS Newhaven Road site 2 CQ 692 614 200 D'orazio, R & Soccol, D. 15 4 20 -Apr- 93 Mawbanna, N.W. TAS Newhaven Road site 3 CQ 706 632 90 D'orazio, R & Socco!, D. 30 4 21 -Apr- 93 Sisters Beach, N.W. TAS Rocky Cape National Park Site 1 CQ 792 684 30 D'orazio, R &Soccol, D. 4 1 21 -Apr- 93 Sisters Beach, N.W. TAS Rocky Cape National Park site 2 CQ 802 679 50 D'orazio, R &Soccol, D. 0 0 17·May- 93 Smithton, N.W. TAS Bass Highway CQ 144 641 40 D'orazio, R & Soccol, D. 22 5 17 ·May- 93 Roger River, N.W. TAS Salmon River Road, Salmon River CQ 199 534 50 D'orazio, R &Soccol, D. 45 5 17·May- 93 Roger River, N.W. TAS Roger River F.R. CQ 331 524 50 D'orazio, R &Soccol, D. 24 6 17·May- 93 Trowutta, N.W. TAS Trowutta Caves S.R. CQ 408 513 190 D'orazio, R &Soccol, D. 18 5 18·May- 93 Balfour, N.W. TAS Balfour Track F.R. CQ 279 439 50 D'orazio, R &Soccol, D. 32 4 18·May- 93 Roger River, N.W. TAS Horton River, Sumac Road. CQ 345 324 190 D'orazio, R. &Soccol, D. 16 2 18·May· 93 Roger River, N.W. TAS Julius River F.R. CQ 345 421 110 D'orazio, R &Soccol, D. 18 3 18·May- 93 Roger River, N.W. TAS Lake Chisholm F.R. CQ 373 444 120 D'orazio, R. &Soccol, D. 12 2 18·May- 93 Roger River, N.W. TAS Horton River CQ 401 251 230 D'orazio, R. &Socco!, D. 21 2 18·May- 93 Roger River, N.W. TAS Wes Beckett F.R. CQ 433 386 180 D'orazio, R &Soccol, D.

19 6 19·May- 93 Marrawah, N.W. TAS West Point CO 024 566 30 D'orazio, R. & Soccol, D. 15 3 19·May· 93 Temma, N.W. TAS Heemskirk Road CO 125 372 100 D'orazio, R. & Soccol, D. 34 4 19·May- 93 Marrawah, N.W. TAS Dismal Swamp Reserve road CO 193 626 60 D'orazio, R. & Socco!, D. 15 2 19·May- 93 Temma, N.W. TAS Blackwater Road CO 227 384 90 D'orazio, R. & Soccol, D. 1 1 31·May· 93 Waratah, N.W. TAS Jasper Hill Track CO 605 058 210 D'orazio, R. & Soccol, D. 20 1 31·May- 93 Waratah, N.W. TAS Mount Cleveland Road CO 618 109 210 D'orazio, R. & Soccol, D. 1 31·May- 93 Waratah, N.W. TAS Wallaby Ck Road CO 721 054 685 D'orazio, R. & Soccol, D. 25 4 31·May- 93 Waratah, N.W. TAS Belmont Road CO 769 174 390 D'orazio, R. & Soccol, D. 15 3 31·May- 93 Hellyer Gorge Reserve, N.W. TAS Hellyer Gorge CO 836 294 310 D'orazio, R. & Soccol, D. 4 1 1 ·Jun- 93 Tullah, N.W. TAS Lower Pieman Road site 1 CP 577 814 230 D'orazio, R. & Soccol, D. 6 3 1 -Jun- 93 Tullah, N.W. TAS Lower Pieman Road site 2 CP 592 809 200 D'orazio, R. & Soccol, D. 33 6 1 -Jun- 93 Tullah, N.W. TAS Lower Pie man Road site 3 CP 623 809 170 D'orazio, R. & Soccol, D. 41 4 1 -Jun- 93 Tullah, N.W. TAS Lower Pieman Road site 4 CP 643 814 170 D'orazio, R. & Soccol, D. 21 5 1 -Jun- 93 Tullah, N.W. TAS Lower Pieman Road site 5 CP 703 785 160 D'orazio, R. & Soccol, D. 33 3 2 -Jun- 93 Corinna, W. TAS S.R. CP 403 875 40 D'orazio, R. & Soccol, D. 37 3 2 -Jun- 93 Corinna, W. TAS Little Hunter Ck CP 445 910 200 D'orazio, R. & Soccol, D. 24 3 2 -Jun· 93 Corinna, W. TAS Timbs Ck CP 450 911 200 D'orazio, R. & Soccol, D. 25 5 15 -Jun- 93 , N.W. TAS Southwell River Road CP 970 989 610 D'orazio, R. & Soccol, D. 34 4 15 -Jun· 93 Cradle Mountain, N.W. TAS Etchells Ck DP 069 981 910 D'orazio, R. & Soccol, D. 15 4 15 -Jun- 93 Cradle Mountain, N.W. TAS Dove River Road DP 169 959 680 D'orazio, R. & Soccol, D. 27 9 16 -Jun- 93 Cradle Mountain, N.W. TAS Murchison Highway S.R. CP 901 976 680 D'orazio, R & Soccol, D. 31 5 16 -Jun· 93 Cradle Mountain, N.W. TAS Cradle Mountain road CP 949 986 670 D'orazio, R & Soccol, D. 14 1 16 -Jun- 93 Cradle Mountain, N.W. TAS Beecroft Road CO 995 034 680 D'orazio, R. & Soccol, D. 4 2 26 -Jul- 93 Zeehan, W. TAS CP 675 598 160 D'orazio, R. & Soccol, D. 9 4 26 -Jul· 93 Zeehan, W. TAS site 1 CP 771 505 380 D'orazio, R & Soccol, D. 3 3 26 -Jul- 93 Zeehan, W. TAS Anthony Road site 2 CP 839 609 530 D'orazio, R. & Soccol, D. 25 4 26 ·Jul· 93 Zeehan, W. TAS Anthony Road site 3 CP 854 671 580 D'orazio, R. & Soccol, D. 21 6 26 -Jul- 93 Zeehan, W. TAS Anthony Road site 4 CP 861 731 400 D'orazio, R & Soccol, D. 21 5 27 -Jul· 93 Zeehan, W. TAS CP 379 689 40 D'orazio, R. & Soccol, D.

20 28 5 27 -Jul- 93 Zeehan, W. TAS Heemskirk Road site 1 CP 523 689 170 D'orazio, R. & Soccol, D. 17 5 27 -Jul- 93 Zeehan, W. TAS Heemskirk Road site 2 CP 568 669 185 D'orazio, R. & Soccol, D. 20 4 27 -Jul- 93 Zeehan, W. TAS Heemskirk Road site 3 CP 601 634 210 D'orazio, R. & Soccol, D. 30 5 27 -Jul- 93 Zeehan, W. TAS Henty Road site 1 CP 620 494 80 D'orazio, R. & Soccol, D. 1 1 27 -Jul- 93 Zeehan, W. TAS Henty Road site 2 CP 630 585 140 D'orazio, R. & Soccol, D. 53 8 28 -Jul- 93 Zeehan, W. TAS Henty Road site 3 CP 561 460 50 D'orazio, R. & Soccol, D. 10 3 27 -Jul- 93 Zeehan, W. TAS Little CP 641 549 140 D'orazio, R. & Soccol, D. 7 1 28 -Jul- 93 Zeehan, W. TAS Road site 1 CP 536 592 225 D'orazio, R. & Soccol, D. 52 8 28 -Jul- 93 Zeehan, W. TAS Trial Harbour Road site 2 CP 593 616 220 D'orazio, R. & Soccol, D. 3 3 9·Aug- 93 Queenstown, W. TAS Lyell Highway site 1 CP 735 324 220 D'orazio, R. & soccor, D. 42 6 9·Aug- 93 Queenstown, W. TAS Lyell Highway site 2 CP 748 336 340 D'orazio, R. & Soccol, D. 20 2 9·Aug- 93 Queenstown, W. TAS Lyell Highway site 3 CP 754 376 300 D'orazio, R. & Soccol, D. 37 4 9·Aug- 93 Queenstown, W. TAS Rd CP 795 456 360 D'orazio, R. & Soccol, D. 18 4 10·Aug- 93 Queenstown, W. TAS Road site 1 CP 853 156 220 D'orazio, R. & Soccol, D. 34 8 10·Aug- 93 Queenstown, W. TAS Kelly Basin Road site 2 CP 860 213 200 D'orazio, R. & Soccol, D. 9 4 10·Aug- 93 Queenstown, W. TAS Kelly Basin Road site 3 CP 900 083 420 D'orazio, R. & Soccol, D. 25 7 10·Aug- 93 Queenstown, W. TAS CP 787 315 35 D'orazio, R. & Soccol, D. 11 5 10·Aug- 93 Queenstown, W. TAS Road site 1 CP 838 306 625 D'orazio, R. & Soccol, D. 14 5 10·Aug- 93 Queenstown, W. TAS Mount Jukes Road site 2 CP 857 255 260 D'orazio, R. & Soccol, D. 11 7 11 ·Aug- 93 Queenstown, W. TAS Lyell Highway CP 895 406 235 D'orazio, R. & Soccol, D. 19 4 11 -Aug- 93 Queenstown, W. TAS Nelson River CP 954 377 330 D'orazio, R. & Soccol, D. 8 2 11-Aug- 93 , Central Plateau, TAS Collingwood River DP 113 314 340 D'orazio, R. & Soccol, D. 5 3 11·Aug- 93 Mt. Arrowsmith, W. TAS DP 188 257 395 D'orazio, R. & Soccol, D. 2 1 11 ·Aug- 93 Derwent Bridge, Central Plateau, TAS Little Navare River DP 303 295 820 D'orazio, R. & Soccol, D. 11 1 11·Aug- 93 Miena, Central Plateau, TAS Tods Corner DP 821 559 1060 D'orazio, R. & Soccol, D. 20 4 11 -Oct- 93 Maydena, S.E. TAS Styx Road site 1 ON 686 592 325 D'orazio, R. & Soccol, D. 38 8 11 -Oct- 93 Maydena, S.E. T AS Styx Road site 2 ON 751 611 340 D'orazio, R. & Soccol, D. 33 4 11 -Oct- 93 Uxbridge, S.E. TAS Styx Road site 3 ON 812 637 250 D'orazio, R. & Soccol, D. 21 7 11 -Oct- 93 Uxbridge, S.E. T AS Styx Road site 4 ON 834 668 240 D'orazio, R. & Soccol, D.

21 24 4 11 -Oct- 93 Uxbridge, S.E. TAS Plenty Valley Road ON 885 549 470 D'orazio, R. & Soccol, D. 17 5 12 -Oct- 93 Mt. Wedge, S.W. TAS Road ON 428 344 260 D'orazio, R. & Soccol, D. 23 6 12 -Oct- 93 Mt. Wedge, S.W. TAS Scotts Peak Road ON 482 413 320 D'orazio, R. & Soccol, D. 1 12 -Oct- 93 Mt. Wedge, S.W. TAS Scotts Peak Road ON 491 526 360 D'orazio, R. & Soccol, D. 15 3 12 -Oct- 93 Maydena, S.E. TAS Scotts Peak Road ON 496 575 570 D'orazio, R. & Soccol, D. 9 3 12 -Oct- 93 Maydena, S.E. T AS Mueller Road ON 532 598 540 D'orazio, R. & Soccol, D. 15 5 12 -Oct- 93 Maydena, S.E. TAS Mueller Road ON 584 606 520 D'orazio, R. & Soccol, D. 24 7 12 -Oct- 93 Maydena, S.E. TAS Mueller Road ON 625 610 520 D'orazio, R. & Soccol, D. 25 4 12 -Oct- 93 Maydena, S.E. TAS Styx Road ON 656 624 540 D'orazio, R. & Soccol, D. 9 2 13 -Oct- 93 Mt. Field National Park, S.E. TAS ON 663 738 1070 D'orazio, R. & Soccol, D. 4 1 13 -Oct- 93 Mt. Field National Park, S.E. TAS ON 694 746 990 D'orazio, R. & Soccol, D. 1 1 13 -Oct- 93 Mt. Field National Park, S.E. T AS ON 733 743 620 D'orazio, R. & Soccol, D. 29 3 13 -Oct- 93 Mt. Field National Park, S.E. TAS ON 751 744 280 D'orazio, R. & Soccol, D. 28 6 13 -Oct- 93 Mt. Field National Park, S.E. TAS ON 764 747 200 D'orazio, R. & Soccol, D. 17 5 16 -Oct- 93 Birch Inlet, W. TAS Birch Inlet eN 753 878 10 Griffith, J. K. & Swiatkowski, P. 10 3 19 -Oct- 93 Avoca, E. TAS Buffalo Brook EP 488 788 360 Kingston, T.J. & D'orazio, R. 21 4 19 -Oct- 93 Avoca, E. TAS Buffalo Brook EP 492 791 320 Kingston, T.J. & D'orazio, R. 27 2 20 -Oct- 93 Epping Forest, N. T AS Mount Joy Road EP 185 724 160 Kingston, T.J. & D'orazio, R. 1 1 9·Nov- 93 Avoca, E. TAS Milford Road EP 467 735 265 D'orazio, R. 5 3 9·Nov- 93 Avoca, E. TAS Milford Road EP 477 744 390 D'orazio, R. 1 1 9·Nov- 93 Avoca, E. TAS Milford Road EP 735 467 265 D'orazio, R. 11 1 23 ·Nov- 93 Blackmans Lagoon, N.E. TAS Blackmans lagoon EO 498 704 20 Kingston, T.J. & Griffith, J. 15 1 23 -Nov- 93 Waterhouse, N.E. TAS Marys Puddle EO 535 784 20 Kingston, T.J. & Griffith, J. 14 3 11 -Jan- 94 Winnaleah, N.E. TAS Banca Road EO 670 595 180 D'orazio, R. & Socco!, D. 8 3 11 -Jan- 94 Winnaleah, N.E. TAS Banca Road EO 684 632 40 D'orazio, R. & Soccol, D. 12 2 11 -Jan- 94 Winnaleah, N.E. TAS Banca Road EO 689 573 110 D'orazio, R. & Soccol, D. 13 2 11 -Jan- 94 Waterhouse, N.E. TAS Waterhouse Road EO 724 693 20 D'orazio, R. & Socco!, D. 6 1 11 -Jan- 94 Waterhouse, N.E. TAS Waterhouse Road EO 838 649 40 D'orazio, R. & Soccol, D. 17 3 11 -Jan- 94 Gladstone, N.E. TAS Browns Bridge Rd EO 871 664 60 D'orazio, R. & Soccol, D.

22 3 1 12 -Jan- 94 Gladstone, N.E. TAS Road EQ 970 636 40 D'orazio. R. & Soccol. D. 13 1 12 -Jan- 94 Gladstone, N.E. TAS Eddystone Road EQ 981 590 40 D'orazio. R. & Soccol, D. 5 1 12 -Jan- 94 Gladstone, N.E. TAS Mt. William National Park FQ 002 726 40 D'orazio. R. & Soccol, D. 2 1 12 -Jan- 94 Gladstone, N.E. TAS Mt. William National Park FQ 015 728 40 D'orazio, R. & Soccol. D. 7 2 12 -Jan- 94 Gladstone, N.E. TAS Mt. William National Park FQ 029 739 10 D'orazio, R. & Soccol, D. 12 2 12 -Jan- 94 Ansons Bay, N.E. TAS Ansons River Reserve FQ 034 541 40 D'orazio, R. & Soccol, D. 1 1 12 -Jan- 94 Ansons Bay, N.E. TAS Eddystone Road FQ 044 582 80 D'orazio, R. & Soccol, D. 3 1 12 -Jan- 94 Ansons Bay, N.E. TAS Eddystone Road FQ 098 598 40 D'orazio, R. & Soccol, D. 3 1 12 -Jan- 94 Ansons Bay, N.E. TAS Eddystone Road FQ 108 602 10 D'orazio, R. & Soccol, D. 3 2 19 -Jan- 94 Upper Castra, N.W. TAS White Rock Road DQ 216 234 470 D'orazio, R. & Socco', D. 27 5 19 -Jan- 94 Upper Castra, N.W. TAS Gaunts Road DQ 228 213 485 D'orazio, R. & Socco' , D. 14 4 19 -Jan- 94 Upper Castra, N.W. TAS Flints Road DQ 228 224 480 D'orazio, R. & Soccol, D. 10 1 19 -Jan- 94 Upper Castra, N.W. TAS Brambles Road DQ 292 276 340 D'orazio, R. & Soccol, D. 22 3 19 -Jan- 94 Upper Castra, N.W. TAS Chilcotts Road DQ 304 274 285 D'orazio, R. & Soccol, D. 4 1 19 -Jan- 94 Upper Castra, N.W. TAS Swamp Road DQ 306 295 200 D'orazio, R. & Soccol, D. 4 1 19 -Jan- 94 Upper Castra, N.W. TAS Chilcotts Road DQ 316 266 240 D'orazio, R. & Soccol, D. 5 2 20 -Jan- 94 Ferndene, N.W. TAS Ferndene DQ 183 377 380 D'orazio, R. & Soccol, D. 13 5 20 -Jan- 94 Penguin, N.W. TAS Penguin DQ 188 445 75 D'orazio, R. & Soccol, D. 10 5 20 -Jan- 94 Penguin, N.W. TAS Penguin DQ 206 416 50 D'orazio, R. & Sacco', D. 1 1 20 -Jan- 94 Penguin, N.W. TAS Penguin DQ 208 458 160 D'orazio, R. & Soccol, D. 2 1 20 -Jan- 94 Penguin, N.W. TAS Mt. Montgomery S.R. DQ 212 451 130 D'orazio, R. & Soccol, D. 15 3 20 -Jan- 94 Penguin, N.W. TAS Dial Road DQ 216 432 145 D'orazio, R. & Soccol, D.

23 What were the outputs of the project?

A high quality comprehensive and representative collection of Tasmanian earthworms, sorted to 'morphospecies·

During the course of the NEGP funded study an additional 340 widely ranging sites were visited specifically to survey earthworms. All sites visited by the author and associates at the Queen Victoria Museum during the period 1990 to 1995 are shown on an outline map of Tasmania (Figure 1). Details for each of the additional 340 sites, including date, grid reference, altitude, the collectors and the number of both specimens and morphospecies retained, are given in Table 4.

Some statistics for the four collection categories previously summarised (Tables 1-4) are presented in Table 5. These include the mean number of specimens collected at sites, mean number of specimens per specimen lot (a specimen lot refers to all the specimens of a given morphospecies at a given site), the mean number of specimens per morphospecies and the mean number of morphospecies per site. (A morphospecies is a readily distinguishable variety that has not yet been subjected to the formal taxonomic process required to determine whether it is a truly distinct species.) Comparison of the summary of collections by the present author (Tables 3 and 4) with those of previous collections (Tables 1 and 2) reveals that the present study was more extensive and intensive, resulting in a 10-fold increase in the number of specimens available for study and a 5-fold increase in the number of sites (Table 5). However, even these figures fail to tell the full story of the improved situation with respect to Tasmanian earthworm collections. Other respects in which the quality of the collections in the present study greatly surpass those of preceding ones include:

• Systematic planning of the sampling programme to ensure the most comprehensive coverage of Tasmania, including Flinders, Maria and Bruny Islands, and to avoid duplication of sampling (Figure 1).

• Application of extensive previous sampling experience led to the formation of an efficient 'search image' for earthworms, and to knowledge of where best to locate specimens and how to extract deep­ burrowing species without damage.

• Strategic collecting, whereby specimens of the same species as already collected were discarded, and the discipline to move on to a new habitat in a timely fashion, prevented col/ections from being dominated by a large number of specimens of one or two common species. (Acknowledgement that this technique was employed requires that the proportions of species in the collections be taken as an approximate, rather than absolute, reflection of relative abundance of species in the field.)

24 Figure 1 Distribution of survey sites by the author and associates 1990-94

25 TABLE 5 Summary of statistics for earthworm collections since 1930

Notes: Includes specimens from targetted collecting of earthworms or ground invertebrates only

1 A "specimen lot" comprises the specimens collected at a single locality that belong to the same species 2 Species not re-collected since 1930 are excluded 3 "Species" in this study refers to a morphospecies, ie a recognisably distinctive variety

collections sites I specimens I specimen species Ispecimens I specimens I specimens I species lots1 I site I lot I species I site

Collections from 1930 -1971 63 396 107 34 6.3 3.7 11.6 1.7 included in Jamieson (1974)2

Collection of Jamieson and Walker 23 399 36 193 17.3 11.1 21.0 1.6 in 1972

Collections by Kingston and 64 951 170 563 14.9 5.6 17.0 2.7 associates at QVM 1990-92

Collections under present study 340 6821 1089 2133 20.1 6.3 32.0 3.2

Combined studies 490 8567 1402

26 • Standardisation of total sampling effort at each site at 2 man-hours throughout the survey, as well as of the techniques described above, ensured that the overall number of specimens and number of species collected at a site provided an accurate reHection of its true relative abundance and diversity. Such relative abundance between sites cannot be inferred for any previous studies of Tasmanian earthworms.

• The quality of georeferencing of the collections, although not obtained through GPS, was nevertheless accurate to within 100m in most cases and thus superior to previous studies. This was achieved through the consistent use of the 1:25 000 map series to identify geographic features and the survey vehicle odometer that was readable to 50m accuracy. In contrast, early collections were referenced (or retrospectively judged) either to the nearest 5 minutes (records in Jamieson 1974) or I minute (Jamieson and Walker collections) of latitude and longitude, representing a fixing accuracy of about 8km and 1.5km respectively. Greater accuracy clearly has benefits in that it allows uncommon species to be re-collected if necessary and the determination of distribution in relation to Reserve boundaries where a species is found to be rare. Such accurate determination of a sampling location has the added benefit that the altitude of the site can be determined from the map with a consistently high level of accuracy, information that will eventually have potential value in defining the ecological envelope occupied by each species.

• The best information available on the chemical fixing and preservation of earthworm specimens was extracted from the literature and used in the survey. Given that preservation is most efficiently conducted in the laboratory setting, the use of moist sphagnum moss in large, well aerated, plastic containers was adopted as a superior method for storage in the field and, in conjunction with a good quality insulated box or 'esky', for transport back to base. This system had a very significant advantage in that if a single damaged worm died and decomposed in the jar the resulting ordure was absorbed by the sphagnum and did not cause the death of the entire batch. '8est practice' fixation of specimens was extended through a new technique, developed by the author, whereby specimens are anaesthetised in 10% ethanol and then suspended on a thread into the formalin fixation bath. This ensures that the specimens are preserved elongated, rather than coiled, and makes later microscopic examination and dissection a more tractable process. Few of the early collections were apparently fixed at all and many are now in poor condition as a consequence, whereas the author's collections would be expected to show negligible deterioration over decades.

The mean number of specimens collected from each site (6.3) was especially low for the 1938 to 1971 collections, reflecting the fact that many of the earlier collections were opportunistic rather than based on survey. This highlights the limitation of the collections upon which Jamieson (1974) was based. The mean

27 figure of 20.1 specimens per site in the present study was achieved through spending 2 man-hours of effort at each site. The results suggest that this collecting effort would be a good benchmark for future earthworm surveys.

The number of morphospecies found at a site varied in the present study from zero to nine (Figure 2), with the great majority of sites (76%) returning from one to four. Nine were found at two sites in the NEGP study and at one site in 1990 by Bob Mesibov. Eight morphospecies were found at seven sites in the present study. The greatest number of (true) species found at a site in the early collecting was 5 by John Hickman at Fern Glade, Burnie (Table 1); while Jamieson and Walker collected more than 2 species from only 2 sites, including their maximum of 7 at one site near Waratah (Table 2). The mean of 3.2 morphospecies per site in this study (Table 5) is double the number of species of the early collectors. This is a result partly due to a greater search effort at each site and partly to the strategic collecting method, looking in all available micro- at each site, together with perhaps a willingness to dig more deeply in the soil than less focused collectors. The majority of the sites at which no native morphospecies were recovered were instead populated by exotic species, notably the European Lumbricidae family. These were all sites that were on the margins of agricultural land or land that was incompletely cleared of native vegetation. They were sampled to check for persisting native species. Two groups of commonly occurring lumbricids, categorised as morphospecies 36 and 44, are included among the sketches and distribution maps in Volume 2. However, no records of species known to be introduced into Tasmania are included in the Tables and summary statistics.

The number of specimens per morphospecies varied greatly in the present study (Figure 3), from ten that were represented by just a single specimen each, to one, Megasco/ex montisarthuri, for which 698 specimens were gathered from 59 sites. The mean number, 32 specimens per site, was much higher in the present study than any of the previous ones, despite the far greater number of species and despite the discarding of multiple specimens of one or two abundant species at many sites. The reason for the high number is the relatively much higher intensity of the sampling; resulting in many morphospecies being collected at multiple sites.

A fourth statistic, the number of sites occupied by each morphospecies varied between one and 59 sites (Figure 4). 50 morphospecies were found only at a single site, while the most wide-ranging species (Megasco/ex montisarthuri again) was found at the maximum number of 59 sites.

A set of sketches of the external secondary sexual characters of about 212 morphospecies

All of the specimens collected were examined and, if not previously encountered, a sketch was drawn of the arrangement of the secondary sexual characters or "genital field". This arrangement, together with features such as size, colour of pigmentation and body proportions were used to sort specimens to the distinct categories referred to as 'morphospecies'. Copies of aI/ of these

28 Figure 2 Frequency distribution of the number of earthworm morphospecies found at each site

100

90

80

70

(J) Q) 60 ::: (J) 0 -... 50 Q) .0 E :::::I s:::: 40

30

20

10

0 0 2 3 4 5 6 7 8 9 number of earthworm morphospecies found at a site

29 Figure 3 Frequency distribution of the number of specimens collected of each morphospecies

50

45 I[ij

40

:{l 35 (.) Q) c.. II) 30 0 ..!: ....c.. o 25 ....E 0 Q; 20 ..c E ~ 15

5

0 ~~~~~~~~~w~~~~~~~~~& "C) ()C) ~C) ~C) ~C) C)C) ~C) ~C) 9:)~ '" ~ ~ ~v~v~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~'l) ~'(;; ~"3 1(5 ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~" ~"V ~" ~'" 10" " "C) ,," ,,~ ,,~ ,,~ ,,~ ~C) rV number of specimens collected

30 Figure 4 Frequency distribution of the number of sites at which morphospecies were found

60 ~------~

50

(/) ~ 40 0 Q) a. (/) 0 ..r:: ...a. o 30 E -...0 Q) .0 § 20 c

10

o f/..t [ / ,1 1( ...' ...... 4 r ...... ] rc.1 t,·",-... r.··A V / J ,...... J ,." =11 r id I// J ...... J ' · ..· ,1 v ..'J l//J VI" l//J V"'J V/J V i" VI] _v ...·J V / ' V..., V/J I 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 26 29 33 34 39 46 59 number of sites

31 sketches are included in Volume II of this report. Although these sketches proved invaluable in sorting specimens to morphospecies, these categories must be seen only as provisional ones requiring confirmation through dissection and description of internal organs.

Distribution maps for the defined morpho-species

Once all of the collected specimens had been sorted to morpho-species and the records collated, the distribution of each was plotted on an outline map of Tasmania using the computer software package IDRISI. A copy of the map for each morpho-species is included in Volume II of this report, alongside the sketch of the same morphospecies. These maps, once revised to take account of changes to morphospecies categories required by taxonomic study, will form the basis of a planned publication "Atlas of Tasmanian Earthworms".

Information on the relationship between earthworms and soil profile

During the course of the present study the author became aware that Mr Mike Laffan, a soil scientist working with Forestry Tasmania, was conducting research into soil profiles in northeast Tasmania. Mr Laffan's observations of the co-occurrence of deep-burrowing earthworms and soil profiles that exhibited much mixing of different sized particles had led him to the hypothesis that earthworms are important, certainly in maintaining, if not in actually forming these profiles. The opportunity was taken to synchronise our studies at some common sites and to attempt to examine quantitatively the relationship between soil profile type and earthworm species composition and abundance. Such a relationship was indeed demonstrated and documented by Laffan and Kingston (1997). A copy of this paper is appended to Volume I of this report.

A new estimate of the number of earthworm morphospecies present in Tasmania

As mentioned earlier in this report, the number of described species known from Tasmania prior to this study was 46. Of these, 26 were recognised among the collections made in the course of the present study. In addition to these named species a further 186 morphospecies were defined on the basis of external characters alone. The dissection and taxonomic study of these varieties was beyond the scope of the NEGP-funded component of the author's studies and thus there remains some uncertainty as to the actual number of species that will eventually be determined from these collections. Undoubtedly in some cases two or more morphospecies will be found to be merely variations of a single true species, equally probably some morphospecies will be found to separate out into two or more distinct species once internal morphology is examined. Currently however, the best possible estimate of the total number of 'species' of Tasmanian native earthworms is 232, made up of the following:

32 • Species previously recorded for the state as included in the catalogue of Jamieson (1974), but not collected in the present study. (approx. 20 taxa)

• Taxa previously recorded for the state and included in the list of Jamieson (1974), and collected again during the present study. (approximately 26 taxa)

• Taxa recorded for the first time during the present study (approximately 186 morphospecies)

The overall estimate for the number of morphospecies calculated from these figures will clearly depend upon how one deals with the 20 "uncollected' taxa of Jamieson (1974). Many of these were described from very few specimens, were only sparsely described and the specimens have not survived. Others have probably been collected in the present study but different preservation techniques may have resulted in apparent differences in external morphology. For these reasons, and to avoid exaggerating the total number of morphospecies, these 20 are assumed to be represented but undetected within the collections accumulated in the current study. On this basis the currently known number of morphospecies is 212.

How many earthworm morphospecies are there in Tasmania?

As already noted the number of described species and new morphospecies combined has been increased during this study from 46 to 212. What proportion of the total number of morphospecies present in Tasmania is this figure likely to represent?

Although the field survey in this study was very extensive, and as comprehensive as it could have been with the resources available, there are clearly some inaccessible parts of Tasmania that were not sampled (Figure 1). Quite extensive gaps between samples exist particularly in the Central Plateau and Western parts of the State. Indeed, apart from the opportunities offered to collect at Pelion and Melaleuca, almost no other samples came from beyond the end of minor roads and forest tracks. Additionally, no sampling was carried out on King Island due to atypically early drying of the soil in spring 1994, when a visit to the Island had been planned.

There are a number of ways of examining the efficacy of the sampling programme. Perhaps the simplest is to consider how many new morphospecies were being found at sites visited late in the collecting period. A plot of the cumulative number of morphospecies collected, against the number of sites visited during the Museum surveys, commencing in August 1990 (Figure 5) reveals that the number of species was still increasing at a perceptible rate even at the end of the survey period. Linear correlation

33 Figure 5 Increasing number of morphospecies found with increasing number of sites sampled

300 ,------,

y =0.57x R2 = 0.93 250

~ 200 ~ QJ ~ c.. (f) 0 ..r:::: ...c.. o 150 E QJ .....> ItS ::::l E 100 ::::l 0

50 -

o !.....­ o 50 100 150 200 250 300 350 400 450 500 cumulative sites

34 between cumulative species and cumulative sites gave a correlation coefficient of 0.93. A least squares best-fit linear plot through the graph's origin is shown for comparison. The data plot commences above the best 'fit line, a feature that is explained by the details of the first six sites in the sequence (Table 3). Three each of these sites were at Weldborough and three at Waratah, widely separated sites with no overlap of species. The mean number of species at these sites was 6.2, more than double the mean for all Museum surveyed sites. Once about 100 sites had been surveyed, the mean number of species collected per site starts to decline, until at about 270 sites it falls below the study average for the first time. After about 350 sites the rate of discovering new species has clearly dropped below the average, however, even in some of the last samples of the study new species were still being collected.

The above finding is perhaps not surprising considering that the survey program moved to unsurveyed regions of the State progressively week by week, the last sites were thus almost equally unrepresented in the collections as was the very first site surveyed. This finding perhaps tells us more about the extent of the distribution of species than it does about the number of them; clearly the fact that additional new morphospecies were found throughout the survey tells us that the range of many of them extends across relatively few consecutive sites. In attempting to use a projection of the relationship between cumulative species and cumulative sites to predict a total number of morphospecies for Tasmania one would clearly have to calculate the extent of unsampled areas of the State. King Island is one readily determined such area, as indeed are extensive remote parts of the central and western regions, equivalent to at least an additional 50 sites.

The calculated slope of the least squares line is 0.57 or, in words, a new morphospecies was identified on average at approximately every second site. In fact what tended to occur when moving into a new survey area was that several new morphospecies would be found at the first site in that area, followed by a rapid decline in the number of additional new morphospecies at subsequent sites within that area. Clearly this generalisation was also affected by the distance between the new area and the nearest previously sampled area. Towards the end of the survey the new areas increasingly lay between areas already sampled, and thus fewer new morphospecies were found. This is consistent with the observed drop of the data trace below the least squares line late in the survey.

An alternative way of examining the collecting data is to add site data in randomised rather than chronological order. This process differs from the above disclJssion in that it says nothing about unsurveyed areas but gives greater insight into the adequecy of the sampling effort within those areas actually sampled. The resulting graph of cumUlative species collected (Figure 6) is clearly curvilinear, implying that a large number of additional sample sites within the same area as previously surveyed would return very few additional morphospecies. The curve appears to be levelling out at about 220 species. On this bais it is reasonable to assume that the predicted maximum number of morphospecies to be found in the area surveyed is of the order of 220.

35 Figure 6 Increasing number of morphospecies found with increasing sites (site order randomised)

250 , ------­ ------­----_ ._­ ,

200 -' ~.,­ !II Q) u Q) ...,..,.,r~ a. !IIo 150 ..c ...a. ,J~ o E Q) .~ > J :;:::; 100 . rt:I ::::l / E ::::l ./ U .// 50 ; ., .~ ., .,. o . o 50 100 150 200 250 300 350 400 450 500 cumulative sites

36 To obtain the best possible estimate for the total number of morphospecies for Tasmania and its offshore islands, 10 additional ones should be allowed for King Island and an additional 20 for the estimated 50 equivalent sites in remote parts of Tasmania that remained unsurveyed. The best estimate possible is thus approximately 250 morphospecies.

How directly are morphospecies likely to translate into true species?

The use of external morphological characters alone to define earthworm varieties in the course of the survey proved to be a very valuable way of rapidly classifying the specimens. Given the large number of forms not found to correspond to any of the species in Jamieson 1974, there was really no choice but to employ this rapid, but admittedly provisional, method., There was an acknowledged expectation that upon dissection some morphospecies would, on the basis of differences in the structure of internal organs, be found to belong to two or more distinct species. Some of the defined morphospecies have distributions (see maps in Volume 2) that strongly hint that species might have been combined. Without wishing to delve too deeply here into the taxonomic work undertaken since the survey, in many cases formal taxonomic study has separated morphospecies in a way that does indeed 'explain' disjunct morphospecies distributions. Similarly some separate morphospecies would be expected to be found to be variations of a single species.

How widely distributed are the individual morphospecies?

Until such time as the translation of morphospecies categories into species is completed there is little point is examining the distributions of individual morphospecies in any detail, or in looking for general patterns. However, the frequency distribution of the number of sites occupied by each of the morphospecies (Figure 4) shows that even after sampling earthworms at over 400 sites across the State, 50 of them, perhaps 20% of the total, had nevertheless been found at only a single locality. Very few sites were situated more than 10 km from the nearest adjacent site which thus implies that these morphospecies had very restricted ranges, in many cases less than 10km in radius. Alternatively these morphospecies might be more widespread than it appears but are simply so uncommon that finding them involves a high degree of chance. Some of these may be associated with specific environmental conditions that are themselves patchy and uncommon, an association that was not recognised during the survey.

At the other extreme, only 10 morphospecies were recorded from more than 20 sites each, implying that very few species have truly extensive distributions. Again this is a picture that could change as the formal taxonomy proceeds if instances are found in which two or more morphospecies are found to be local variations of a single species. With respect to the taxonomic study completed to date this has not however been found to be a common occurrence.

37 What conclusions can be drawn concerning the conservation status of species

Any true species found at fewer than 4 distinct sites within Tasmania would require further, more detailed survey of its distribution and habitat requirements. 139 of the 212 morphospecies, or 65% currently fall into this category.

Species found at just one or two sites would require this work to be done as a matter of high priority. There are 88 morphospecies (42%) that fall into this group at present. Once confirmed through formal taxonomy, species that remain in this category will be the subject of further investigation and assessment of their distribution, abundance and the current and prospective level of threat to their survival. Where appropriate the species will be proposed for inclusion in the formal list of Tasmanian invertebrates under the State's Threatened Species Protection Act 1995.

What recommendations for future work emerge from this project?

The major recommendations for future work following on from this study include:

• Rigorous taxonomic study of all morphospecies defined in this study through examination of all specimens attributed to each morphospecies.

• Supplementary collecting, on an opportunistic basis, in those remote areas of Tasmania not yet able to be visited, as well as on King Island.

• Further more intensive survey in the areas surrounding each of the species that were recorded from only one or two sites, in order to better define the distribution of those species. This survey should also determine the habitat preference of these species to facilitate the next recommendation

• Each species found to be rare, or to occupy a restricted geographical range, should be assessed for listing under the Threatened Species Protection Act 1995 and inclusion in the Threatened fauna manual for production forests in Tasmania (Jackson and Taylor, undated).

38 Postscript: a summary of progress 1994-1999

Since completion of earthworm survey under NEGP funding in 1994 significant additional funds from a variety of sources have been secured and deployed, mostly in support of taxonomic study of the specimens accumulated in the survey. These funds have included:

$61 000 from the Plomley Foundation to fund Dr Rob Blakemore to conduct taxonomic studies of the collections, jointly with the author, between 1996 and 1998.

$5 000 from the Tasmanian Parks and Wildlife Survey to fund a field survey and taxonomic studies of the earthworms around the shores of Lake Pedder in 199

$28 000 from the Tasmanian component of the Regional Forest Agreement process in support of Dr Blakemore's collaboration with the author on earthworm taxonomy.

$65 000 worth of support from the Australian Antarctic Division for studies of earthworms on Macquarie Island.

$33 000 from the Australian Biological Resources Study (ABRS) in support of earthworm taxonomy in 1999/2000

At the time of writing the taxonomic studies are continuing. Detailed descriptions of a total of approximately 180 species of Tasmanian earthworms have been completed, including both new species as well as augmented redescriptions of pre-existing species. One paper has been published (Blakemore and Kingston 1998, Appendix 2) while another has been accepted for publication, subject to only minor revision. By the time the funds listed above have been fully expended it is anticipated that all morphospecies distinguished during the current study will have been examined and all valid species described.

39 References

Blakemore, R.J. and Kingston, T.J. (1997) Opisthogastric earthworms (Megasclecidae: ) and allied forms in north-western Tasmania. Journal of Natural History, 31: 1683-1708

Jamieson. B.G.M. (1974) The indigenous earthworms (O/igochaeta: Megascolecidae) of Tasmania. Bulletin of the British Museum (Natural History) Zoology 26, 201-328.

Kingston, T.J. (1989) Aporrectodea caliginosa and Lumbricus rubel/us populations under irrigated and dryland pastrures in northern Tasmania. In 'Proceedings of the 5th Australasian Conference on Grassland Invertebrate Ecology' Melbourne 1989, pp 199-205.

Kingston, T.J. and Temple-Smith, M.G. (1989) Earthworm populations under Tasmanian pastureland. In 'Proceedings of the 5th Australasian Conference on Grassland Invertebrate Ecology' Melbourne 1989, pp. 192-199.

Laffan, M.D. and Kingston, T.J. (1997) Earthworms in some Tasmanian forest soils in relation to bioturbation and soil texture profile. Australian Journal of Soil Science, 35:1231-43.

Lobry de Bruyn, L.A. and Kingston, T.J. (1997) Effects of summer irrigation and trampling in dairy pastures on soil physical properties and earthworm number and species composition. Australian Journal ofAgricultural Research 48: 1059-79

Temple-Smith, M.G., Kingston, T.J., Furlonge, T.L., and Garnsey, R.B. (1993) The effect of the introduction of earthworms Aporrectodea caliginosa and Aporrectodea longa on pasture production in Tasmania. In Proceedings of the 7th Australian Agronomy Conference "Farming from Paddock to Plate", Agronomy Society of Australia p. 373

Acknowledgements

The superb quality of the earthworm col/ection and associated database built up during this study is a reflection of the total commitment to the project made by the Project Officer Rob D'Orazio. Queen Victoria Museum Research Officer Ms Louise McGowan assisted the project with databasing of information as well as in the field. The project was greatly enhanced by involvement in fieldwork by Mark Cooper under the Commonwealth Government JOBSKILL programme and by a sequence of volunteers including Amanda Mitchell (4 days), Mark Gittus (3 days) and Daniel Soccol (25 days). The production of the maps was made possible through enthusiastic instruction in the use of IDRISI by Glyn Roberts of Howrah. Margot Kingston was selfless in her support of the project, frequently seeing family weekends excursions merge into collecting trips and her kitchen into an earthworm morgue.

40 Appendix 1 Laffan, M.D. and Kingston, T.J. (1997)

Aust. J. Soil Res., 1997, 35, 1231-43

Earthworms in some Tasmanian forest soils in relation to bioturbation and soil texture profile

M. D. LaJjanA and T. J. KingstonB

A Forestry Tasmania, Launceston. Present address: Forest Practices Board, PO Box 180, Kings Meadows, Tas. 7250, Australia. B Queen Victoria Museum, Wellington Street, Launceston, Tas. 7250, Australia.

Abstract Soil properties and earthworm population density were examined for 5 forest soils derived from Silurian-Devonian sandstones (Mathinna Beds) in north-eastern Tasmania. The soils occur along gradients of altitude, rainfall, and forest type; they include 2 with texture-contrast and 3 with gradational soil profile types. The density and biomass of the most abundant earthworm species Megascolex montisarthuri, and of all earthworm species combined, were found to be greater in gradational than in texture-contrast soils. A greater proportion of the earthworms in gradational soils than in texture-contrast soils was found to occur at soil depths exceeding 10 cm. The contrast was most pronounced between the 2 texture-contrast soils and the single gradational soil that occur under dry eucalypt forest. This paper explores the hypothesis that bioturbation of surface and subsurface layers by earthworms is an important mixing process that in gradational soils outweighs the counter tendency for soil particles to sort and thus form texture-contrast profiles.

Additional keywords: soil fauna.

Introduction Soil mapping in north-eastern Tasmanian forests during 1990 by Laffan et al. (1995) differentiated a range of divergent soils formed on Silurian-Devonian sandstone (Mathinna Beds). Of particular interest were 5 soils, including 3 gradational and 2 duplex (texture-contrast) profile forms (Northcote 1979), occurring under dry eucalypt forests and wet forests. These soils comprise the Retreat, Wattley, Piper, Maweena, and Sideling soil profile classes characterised and mapped on the Pipers 1: 100000 scale topographic sheet (41 0 S, 1470 E) (Laffan et al. 1995). The major factors considered responsible for the development of texture-contrast soils have been reviewed by Chittleborough (1992); they include sedimentolOgical layering, clay formation, clay weathering, clay iIluviation and alluviation, and bioturbation. The author noted that bioturbation by soil fauna could cause homogenisation of profiles and thus prevent strong textural differentiation. Where the processes that tend to sort soil particles predominate over long periods, texture contrast profiles may result, whereas when sorting processes are negated by periodic or continuous soil mixing, gradational soils may be sustained. The roles of termites and ants in soil modification, including effects on soil texture profiles, have been reviewed by Lobry de Bruyn and Conacher (1990). Their review showed that there is evidence both for and against texture differentiation

0004-9573/97/061231$05.00 1232 M. D. Laffan and T. J. Kingston

by ants and termites. Research into the role of earthworms in pedogenesis has been reviewed by Lee (1985). More recently, in a study in south-western Ivory Coast, Nooren et al. (1995) proposed that the selective erosion of surface worm casts by splash and overland flow was a significant contributor to the formation of sandy topsoils under forest. A similar process of winnowing of finer soil particles by water and wind erosion of faunal casts and mounds has been proposed as a global mechanism for the development of texture-contrast soils in materials that can be sorted (Paton et al 1995). Conversely, soil mixing by fauna that do not form casts or mounds on the soil surface can counteract the formation of texture-contrast profiles (Humphreys et al. 1996). Field characterisation of the 5 Tasmanian forest soils suggested there may be marked differences in the levels of earthworm activity between soils; those with gradational profiles were observed to contain more subsurface earthworm burrows and casts than those with texture-contrast soils. Surface casts were not observed on any of these soils. Ant mounds occurred sporadically on the surface of the 3 soils under dry eucalypt forest, but there was no evidence that ant mounding was more prevalent on anyone soil. Excavation of ant nests revealed that their galleries generally did not extend below the Al horizon; ant activity is thus unlikely to have caused textural differentiation or mixing of subsurface horizons. Examination of soil morphology and associated site characteristics led the authors to the hypothesis that differing levels of earthworm activity and hence bioturbation, in both surface and subsoil layers, could be a major causal factor in the development of either gradational or texture-contrast profiles. In order to seek support for this hypothesis, the densities of native earthworms, the major potential agents of faunal bioturbation present in these soils, were measured for the 5 soils and were analysed in relation to the 2 contrasting profile types.

Study sites Parent material and topography The parent materials of all 5 soils are Silurian-Devonian sandstone of the Mathinna Group of north-eastern Tasmania (Tasmania Department of Mines 1965). Retreat, Wattley, Piper, and Maweena soils occur on undulating (2°~6°) and rolling (6°-1r) slopes of hills at altitudes of 60~340 m. Parent materials of these soils are mainly strongly weathered. Sideling soils occur on rolling slopes of mountains at altitudes of 480-720 m; their parent materials are less weathered, reflecting the strong impress of periglacial and other colluvial slope processes during the Quaternary period. Retreat soils are predominant on better drained sites, generally on crests and upper slopes of hills, whereas Wattley and Piper soils are more commonly found on less well drained sites such as lower hill slopes and drainage lines. However, Piper soils have been recognised on all segments of hilly landforms apart from narrow rocky ridge crests. In areas of low relief, Wattley soils often occur on lower lying slopes.

Climate and 1Jeyd.tition Mean annual rain:faIl ranges from 800 to 1500 mm and the native vegetation varies from dIy eucalypt forest to wet eucalypt forest and mixed forest of eucalypt ..~ f Table 1. Summary of site and profile data 8 Landform Native vegetationl Texture Drainage Soil classification '" mean annual rainfall profile class AustralianA PPFB Soil ~ TaxonomyC o·C" ",. Retreat ...:: Midslopes, Shrubby Loamy sands over clays Moderately well Bleached, brown Dy5·4J Hapludult g­ upperslopes, forest; drained kurosol ""o· =:: crests of hills 850-1000 mm WatUey Lower slopes of Heathy dry eucalypt Loamy sands over sandy Imperfectly Bleached- Dy5'4l Hapludult hills, crests of forest; clay loams and clays drained brown low rises 850-950 mm chromosol Piper Lower slopes of Sedgy dry eucalypt Fine sandy loams over Imperfectly Acidic-Mottled, Gn3·91 Hapludult drainage forest; sandy clay loarns over drained yellow dermosol lines 80D-IOOO mm sandy clays Maweena All facets of Wet eucalypt forest; Fine sandy loams over Moderately well Acidic-Mottled, Gn3·7l Hapludult hillslopes 100D-1400 mm loams and drained brown dermosol medium clays Sideling Rolling Mixed forest; Clay loams over Well drained Acidic, brown Gn4·3l Dystrochrept mountain slopes 1200-1500 mm light clays dermosol AIsbell (1996). BPrinciple Profile Form, Northcote (1979). cSoil Survey Staff (1992).

l­ t..:> V> V> 1234 M. D. Laffan and T. J. Kingston

canopy over a rainforest understorey. Details of soil and vegetation characteristics for each soil type are provided in Table 1. The groundcover occurring on Piper soils incorporates common to abundant sedges, indicating that surface and!or subsurface soils remain moist for more extended periods than is the case for the other soils. Table 2. Bulk density and particle-size analysis for representative soils Horizon Depth Bulk density SandA SiltB ClayC 3 (ern) (g/cm ) (%) (%) (%) Retreat Al 0-12 1·1 80 12 8 A2 13-24 1·3 84 12 4 A2e 24-35 1·6 88 8 4 Bit 35-41 1·6 56 20 24 B2t 41-83 1·5 24 16 60

Wattley Al 0-18 1·2 82 12 6 A21e 18-30 1·5 84 12 4 A22e 30-36 1·7 84 12 4 B2lt 36-56 1·4 63 12 25 B22t 56-87 1·5 48 12 40 B23g 87-95 n.d. 48 16 36

Piper Al 0-13 1·2 70 16 14 Bit 13-29 1·6 60 16 24 B2t 29-87 1·5 44 16 40 B3gt 87-95 1·5 56 16 28

Maweena Al 0-14 1·1 66 16 18 AB 14-26 1·4 59 20 21 B2lt 26--36 1·8 56 20 24 B22t 36-58 1·6 60 20 20 B23t 58-80 1·6 60 16 24

Sideling Al 0-9 0·8 32 40 28 B21 9--35 1·3 32 44 24 B22 35-70 1·1 28 48 24 n.d., not determined. A2·0-0·02 mm. BO·02-0·002 mm. c<0·002 mm.

Soil properties Soil physical and chemical properties for the 5 soils are given in Table 2 (bulk density and particle-size) and Table 3 (selected chemical characteristics). Retreat and Wattley soils (Fig. 1) are moderately well drained and imperfectly drained, respectively. Both soils have texture-contrast profiles characterised by dark grey sandy loam Al horizons and bleached loamy sand or sand A2e horizons overlying yellowish brown, mottled clayey B2t horizons typically at a depth of about 30 cm Earthworms and bioturbation 1235

in Retreat soils and 40 cm in Wattley soils. Earthworm burrows and casts are moderately abundant in Al and A2 horizons, but little evidence of earthworm activity was recorded in underlying subsoils.

Table 3. Selected chemistry Cor representative soils Horizon Depth pH Total NA OCB Total pC Exch. Ca, ECECE (cm) (%) (%) (%) (jJ.g/g) Mg, K, NaD (cmol/kg) (cmol/kg) Retreat Al 0-13 4·7 0·12 1·1 55 3·84 5·0 A2 13-24 4·8 0·06 1·5 32 n.d. n.d. A2e 24-35 5·1 0·03 0·6 18 n.d. n.d. Blt 35-41 4·9 0·03 0·6 42 n.d. n.d. B2t 41-83 5·3 0·03 0·5 89 1·21 11·0 BC 83-95 5·4 n.d. 0·4 95 n.d. n.d. Wa.ttley Al 0-18 4·8 0·06 1·7 42 1·59 3·0 A21e 18-30 5·1 0·03 0·4 19 n.d. n.d. A22e 30-36 5·6 0·02 0·2 11 n.d. n.d. B2lt 36-56 5·8 0·04 1·0 58 n.d. n.d. B22t 56-87 5·9 0·03 0·3 76 2·51 5·0 B23t 87-90+ 5·9 0·02 0·2 74 n.d. n.d. Piper Al 0-13 4·7 0·08 1·2 52 2·01 4·0 Bit 13-29 4·9 0·03 0·4 47 n.d. n.d. B2t 29-87 5·2 0·03 0·3 76 1·23 13·0 B3gt 87-90+ 5·2 0·02 0·1 113 n.d. n.d. Maweena Al 0-14 5·2 0·19 2·4 179 4·90 6·0 AB 14-26 4·9 0·06 0·9 104 n.d. n.d. B2lt 26-36 4·9 0·05 0·5 92 n.d. n.d. B22t 36-58 4·8 0·04 0·5 142 1·84 9·0 B23t 58-80 4·8 0·05 0·6 202 n.d. n.d. Sideling Al 0-9 4·2 0·37 5·6 359 1·59 7·0 B21 9-35 4·7 0·15 2·9 295 n.d. n.d. B22 35-70 4·9 0·12 1·9 310 0·37 3·0 n.d., not determined. ADetermined using Kjeldahl method (Rayment and Higginson 1992). BOrganic carbon, determined by Walkley and Black (1934) colorimetric method. CDetermined by perchloric acid (Olsen and Sommers 1976). DTotai exchangeable Ca, Mg, K, Na determined using 1 M NH4Cl at pH 7·0. EEffective cation exchange capacity (exchangeable cations+exchangeable acidity).

Piper soils (Fig. 2) are also imperfectly drained, but they have gradational texture profiles characterised by dark grey sandy loam Al horizons overlying olive brown sandy clay loam Bl horizons with distinct grey mottles and many prominent very dark grey infilled earthworm burrows and casts (Fig. 3). Deeper subsoil layers include yellowish brown and greyish fine sandy clays and clay loams with common or few earthworm burrows and casts to a depth of at least 70 cm. In many profiles, earthworm burrows and dark grey casts occur to depths below 90 cm. 1236 M. D. Laffan and T. J. Kingston

Fig. 1. Wattley soil profile to 1 m showing its texture-contrast character.

Maweena and Sideling soils are moderately well drained and well drained, respectively, and typically have gradational texture profiles, although some Sideling soils have uniform profiles. Maweena soils typically have dark brown sandy loam or sandy clay loam Al horizons overlying yellowish brown clay loams and light clays with common distinct clay skins. Sideling soils are characterised by dark greyish brown clay loam Al horizons overlying brown light clay or clay loam subsoils generally with common to many rock fragments. Humic litter layers often occur on the Al horizons of both soils. Numerous earthworm burrows and casts occur to depths of at least 70 cm in both soils.

Earthworm populations Extensive information on the native earthworms of north-eastern Tasmania has been gathered during the course of an ongoing survey commenced in 1992 (T. J. Kingston unpublished data). A minority of the earthworm species recovered have been identified by using species descriptions in Jamieson (1974); the collections were, however, found to contain several undescribed species. Earthworms and bioturbation 1237

Fig. 2. Piper soil profile to 0·8 m showing its gradational character.

Earthworms found within the current study area include species belonging to the 3 main ecological types defined by Lee (1985). Earthworms belonging to the first category, those confined to the litter horizon, were present but are not considered in this study due to their absence from the mineral component of the soil profile. The second category, species that occupy the upper levels of the profile but do not form open burrows, was represented by 2 species: J'vJegascolex montisarthuri and an undescribed species of Vesiculodrilu8 . During the earthworm survey, M. montisarthuri had been found to be widespread and abundant in north-eastern Tasmania (T. J. Kingston unpublished data). The same species was found to be the predominant one for all soils within the present study. During winter, when the soil is wet, members of this species were found to be active in the upper portion of the soil profile. In aJJ 5 soils, the great majority of individuals were found in the upper 20 cm of the profile; however, in the gradational soils, but not in the texture-contrast soils, a minority of earthworms were encountered at depths to 40 cm. In summer, when t he water content of the surface soil declines, the absence of M. montisarthuri individuals in the upper part of the profile has been noted; it appears that they burrow more deeply at this time 1238 M. D. Laffan and T. J. Kingston

Fig. 3. Piper soil subsoil clod (from 40 to 50 cm depth) showing darkly coloured earthworm casts (scale bar 1 cm).

of year, presumably to avoid the desiccating conditions that prevail in the uppermost part of the profile. The second ' topsoil' species, Vesiculodrilus sp., was encountered relatively infrequently compared with M. montisarthuri; nothing is known of the ecology of this species. The third category of earthworm lifestyle, occupation of open burrows that extend deeply into the subsoil, is represented, , in some parts only of the study area, by Tasmania's largest known earthworm species, Pinguidrilus tasmanianus. Mature individuals of this species achieve 40 cm in length, when extended. When present, members of the species occupy semi-permanent burrows that commonly reach a depth of 80 cm. Although the density of both earthworms and active burrows is always low, many unoccupied and infilled burrows were also detected. Detection was facilitated by the presence in the burrows of darker more highly organic soil translocated from the Ai horizons.

Methods Field investigation of the earthworm populations of the 5 soils was conducted during the winter and spring of 1993. For each soil class , 3 representative but widely spaced (> 1 km apart) localities were selected for the study of earthworm composition and density. At each site, a 100-m transect was laid out and .6 points were located randomly along its length. A 0·25-m2 metal frame having 10-cm-high vertical sides was pushed into the ground at each point and the soil was excavated from within the quadrat in 2 depth intervals: 0-10 cm and >10 cm. Excavation continued until no further earthworms or burrows were evident; the depth at which this occurred .varied between about 20 and 80 crn below the surface. Soil excavated from each level was placed onto plastic sheeting, manually broken up, and sifted for earthworms and other soil fauna. All invertebrates found, including severed fragments , were preserved by placing them into 70% ethanol. In the laboratory, earthworms from each site were sorted to species, placed on absorbent paper to remove excess ethanol, ~..,

fo

Table 4. Earthworm population densities (number/m2), by soil class and depth ~ ?5 Within columns, means followed by the same letter are not significantly different at P == 0·05 0­ c­ Soil Megascolex montisarthuri Pinguidrilus tasmanianus Vesiculodrilus sp. All species Total 5'.... 0-10 em >lOcm Combined Q-1O em >lOcm Q-lOcm >lOcm 0-10 em >lOcm Combined r:::... [ Wattley 21·3b 1·1b 22·4bc 0 0 0 0 21·3e l·lb 22·4c o· Retreat 38·2b 1·3b 39·5be 0 0 0 0 38·2be 1·3b 39·5be ;:l Piper 55·1b 14·9a 70·0ab 0 0 2·2 0·2 57·3b 15·la 72·4b Maweena 97·1a 8·9ab 106·0a 0·2 0·2 4-9 0·9 102·2a lO·Oab 112·2a Sideling 36-4b 14-4a 50-8bc 0 0-4 0 0 36-4bc 14-8a 51-3be Overall P <0-01 <0-05 <0-05 <0-01 <0-05 <0·01

Table 5. Earthworm biomass (g/m2), by soil class and depth Within columns, means followed by the same letter are not significantly different at PO·05 Soil Megasco/ex montisarthuri Pinguidrilus tasmanianus Vesiculodrilus sp. All species Total 0-10 em >10cm Combined Q-1O em >lOcm Q-lOem >10 em Q-1O em >10 em Combined Wattley 12·7e 1·3b l4-0e 0 0 0 0 12·7d 1·3b 14·0b Retreat 19-9bc 'O·9b 20-8be 0 0 0 0 19·ged 0·9b 20·8b Piper 50·9a 20-8a 71·7a 0 0 4·4 0-5 55-3ab 21·3ab 76·6a Maweena 62·9a 8·Bab 71·7a 0·9 2·5 9·1 2·0 72·8a 13·3ab 86·0a Sideling 38·4ab 16·0a 54·4ab 0 18·7 0 0 38·4bc 34·7a 73-la Overall P <0·01 <0-05 <0·01 <0-001 <0·05 <0·01

..... to.:> ~ 1240 M. D. Laffan and T. J. Kingston

and weighed. Earthworms in each sample were counted; where there were severed pieces, only the 'heads' were scored. Invertebrates other than earthworms, found mostly in the leaf litter layer, were collected from all sites. Collections included millipedes, centipedes, spiders, beetles, and the larvae of beetles, flies, and moths. Few of these were as large as an individual earthworm and none were found at depths below 25 cm. Together, these invertebrates were responsible for an insignificant proportion of total invertebrate biomass and hence of faunal activity in the soil. As with small litter-dwelling earthworms, these other invertebrates are not considered further in this paper. Results The mean densities of each of the 3 soil-dwelling earthworm species, according to soil class and the 2 depth intervals, are presented in Table 4 (earthworm density) and Table 5 (earthworm biomass). Three trends are apparent from these results. Firstly, for all soil classes, total earthworm biomass was 2·8 times greater at the surface (mean 39·8 gjm2) than below a depth of 10 cm (mean 14·3 gjm2 ). Secondly, for the 2 soil depths combined, total earthworm biomass 2 for the gradational soils (78·6 gjm )) was 4·5 times greater than that for the texture-contrast soils (17·4 gjm2 ). Thirdly, an overall dominance of the results by the one species, M. montisarthuri (86% of total earthworm biomass), is also apparent. Earthworm density and biomass data were log-transformed, and both untrans­ formed and transformed data were examined for normality, by calculation of the Shapiro-Wilks W statistic. For both density and biomass of M. montisarthuri and for the 3 earthworm species combined, log transformation of the data decreased the departure from normality to a non-significant level and thus log-transformed data were employed in all statistical analyses. For the 2 less-abundant species, the distributions of both untransformed and transformed data were far removed from normality and so parametric statistical techniques were unavailable for use in these cases. The transformed data for M. montisarthuri and for all species combined were analysed according to soil profile type and soil depth, using 2-way analysis of variance. A third factor, soil profile type x soil depth interaction, was also included; the results of these analyses are given in Table 6. Soil depth, soil profile type, and the interaction between these were all found to be significant factors in determining earthworm density and biomass in the samples. It is notable that the addition of data for the 2 less-common species to that of M. montisarthuri resulted in only minor changes in the values of the F-ratio. Examination of the coefficient of determination (R2) revealed this to be consistently high at 0·83, whether earthworm density or biomass, or whether M. montisarthuri or combined species were considered. Interpretation ofthe results for soil depth and soil profile type was straightforward; at the time of the study earthworm density and biomass were greater in the upper 10 cm of the soil profile than below this depth. Earthworm density and biomass were also higher in gradational soils than texture-contrast soils. The significant interaction between soil depth and soil profile type may be interpreted as a non-uniform effect of depth between the 2 soil profile types; a greater proportion of earthworm density and biomass was recovered from below 10 cm depth in gradational soils than in texture-contrast soils. Earthworms and bioturbation 1241

Table 6. Result of two-way ANOVA for M. montisarthuri and for all species combined Factor log(earthworm density) log(earthworm biomass) d.f. F-ratio P d.f. F-ratio P Megascolex montisarlhuri Depth 23·1 <0·0001 11·1 0·0003 Soil profile type 8·3 0·0079 17·8 0·0003 Soil profile type x depth 5·5 0·026 4·3 0·047 All species Depth 23·6 <0·0001 17·2 0·0003 Soil profile type 8·8 0-0065 17·2 0·0003 Soil profile type X depth 5·6 0·025 5·6 0·026

While these results demonstrate the influence of soil profile type and soil depth on earthworm density and biomass, they do not distinguish between earthworm density and biomass in the 5 individual soil classes. In order to augment the results in relation to this aspect, univariate analysis of density and biomass by soil class was conducted for M. montisarthuri and for combined earthworm species, and comparisons were carried out between all soil pairs, using Student's t~test. The results, incorporated in Tables 4 and 5, provide insight into which of the 5 individual soil types made the greatest contribution to the results of the 2~way analysis based on the 2 profile types only. Discussion Earthworm density and biomass The sampling methods employed produced estimates of the density and biomass of earthworms that varied in precision, according to species, as determined by calculation of the coefficient of variation (CV). Sampling was more precise for the abundant and widespread M. montisarthuri [CV(density) = 96%, CV(biomass) = 103%], while the size of the areas excavated was less efficient in its estimation of the large and locally occurring species, P. tasmanianus [CV(density) 237%, CV(biomass) 319%)]. The latter species was found only in Maweena and Sideling soils; in each case only 2 individuals were found in the soil excavated from 18 holes. These 4 earthworms imposed a considerable impact on the overall results for biomass, both in terms of the mean values for the sites at which they were found, and in terms of overall variability. For example, for the Sideling soil, inclusion of the 2 P. tasmanianus individuals increased the overall number of earthworms at this site by <1%, but increased the biomass by 34%. Simultaneously, the associated coefficient of variation increased from 23% to 52%. Future studies investigating the impact of large, deeply burrowing earthworms upon soil profile characteristics will need to overcome this problem through the (excessively labour-intensive) process of excavating holes of much larger surface area. Earthworms and the soil texture profile The results show that at the time of the study, in winter, when soil water content was high, there were clear differences between soils with texture-contrast and gradational profiles with respect to the density, biomass, and distribution by depth of earthworms. The proportions of total earthworm bioma..<;s extracted 1242 M. I). !."ffall a/ld T . .1. Kingston

from each of the 2 soil depths vary considerably between soil classes. For the texture-contrast soils, the proportion of earthworm biomass found below 10 cm depth was 9% for Wattley and 4% for Retreat; in contrast, for the gradational soils the equivalent values were 28% (Piper), 16% (Maweena), and 47% (Sideling). It has already been acknowledged above that the earthworm biomass for the Sideling soil was enhanced by the presence of 2 P. tasmanianus. However, even if this species is ignored, the proportion found below 10 cm remains high at 29%. These differences (statistically significant between soil profile types) are consistent with the observation that the gradational soils all have numerous earthworm burrows and casts in their subsoils to depths of at least 70 cm, whereas relatively few burrows and casts were found in subsurface horizons (A2e) and subsoils of texture-contrast soils. The results of the earthworm study, together with profile morphology, strongly suggest that bioturbation by earthworms may be a major factor in the development of gradational texture profiles. Of particular interest are the differences in earthworm biomass and depth of activity of the (texture-contrast) Retreat and Wattley soils as opposed to the (gradational) Piper soil. These soils have similar parent material and climate and all support dry eucalypt forests of similar species composition. The soils are assumed to be of similar age. Variation in the water status of the upper part of the profiles appears to be the main soil feature affecting earthworm populations and distribution, especially since other properties such as bulk density and organic carbon concentrations are relatively uniform (Tables 2 and 3). We infer that upper subsoils of Piper soils remain moist for longer periods and thus provide a habitat for earthworms which is more favourable than the drier subsurface layers in Retreat and Wattley soils. Of course earthworm density and biomass are more likely to have decreased during the formation of the A2e horizons (because of increasing sand content and droughtiness) than to have been stable throughout the development of these subsurface layers. However, we surmise that earthworm biomass would have been relatively low in these soils prior to development of distinct A2e horizons. Examination of the various processes able to influence the development of texture-contrast profiles (Chittleborough 1992) indicates that bioturbation is the most obvious factor which is significantly different between the 3 soils. The impacts of the other factors are difficult to ascertain but it is unlikely that they are different between the soils examined in this study. There is no evidence of sedimentological layering or any lack of uniformity across the soil classes with respect to physical processes, such as forest windthrow, which can cause substantial mixing of surface and subsurface soil layers and thus counteract the development of texture-contrast profiles. Indeed, windthrow would be expected to be more prevalent in Retreat and Wattley soils because of shallower effective rooting depth. The occurrence of distinct clay skins in all 3 soils is taken as evidence of clay illuviation, but we are unable to comment on possible differences in rates of clay translocation. The greater density and biomass ofearthworms in both topsoils and upper subsoils of Piper soils compared with Retreat and Wattley soils is assumed to represent evidence of faunal bioturbation. We suggest that bioturbation by earthworms in the upper layers of Piper soils is a major factor responsible for the development of gradational texture profiles. The continual mixing of the mineral soil fraction Earthworms and bioturbation 1243

between surface and subsoil layers counteracts processes such as clay translocation which favour the development of texture-contrast profiles. Conversely, in Retreat and WattIey soils, the absence of earthworm bioturbation in surface and subsurface layers permits the texture-contrast condition to prevail. Although the results of this study have revealed a distinct relationship between soil profile type and the biomass of earthworms present, it is acknowledged that evidence of causation is circumstantial. The data presented in this study do, however, support the global bioturbation model proposed by Humphreys et al. (1996) to explain the development and maintenance of texture-contrast and gradational or uniform-textured soils. Acknowledgments Laboratory analyses of soils were carried out as part of the forest soils mapping program for the Pipers map-sheet. The authors are grateful to members of the Queen Victoria Museum earthworm survey team, particularly Rob D'Orazio, Greg Finnigan, and Daniel Soccol, for assistance in the field. References ChittIeborough, D. J. (1992). Formation and pedology of duplex soils. Australian Journal of Experimental 32, 815-25. Humphreys, G. S., Mitchell, P. B., and Paton, T. R. (1996). Bioturbation and soil formation: towards a new paradigm. In 'Proceedings Australian and National Soils Conference'. Melbourne, 1-4 July 1996. Isbell, R. F. (1996). 'The Australian Soil Classification.' Australian Soil and Land Survey Handbook Series. Vol. 4. (CSIRO Publishing: Collingwood, Vic.) Jamieson B. G. M. (1974). The indigenous earthworms (Oligochaeta: Megascolecidae) of Tasmania. Bulletin of the British Museum (Natural History) Zoology 26, 201-328. Laifan, M. D., Grant, J. C., and Hill, R. B. (1995). Soils of Tasmanian State Forests. 1. Pipers sheet. Soils Bulletin No.1, Forestry Tasmania, Hobart. Lee, K. E. (1985). 'Earthworms: their Ecology and Relationships with Soils and Land Use.' (Academic Press: London.) Labry de Bruyn, 1. A., and Conacher, A. J. (1990). The role of termites and ants in soil modification: A review. Australian Journal of Soil Research 28, 55-93. Nooren, C. A. M., van Breeman, N., Stoorvogel, J. J., and Jongmans, A. G. (1995). The role of earthworms in the formation of sandy surface soils in a tropical forest in Ivory Coast. Geoderma 65, 135-48. Northcote, K. H. (1979). 'A Factual Key for the Recognition of Australian Soils.' (Rellim Technical Publications: Glenside, S. Aust.) Olsen, S. R., and Sommers, L. E. (1976). Phosphorus. In 'Methods of Soil Analysis'. (Eds A. H. Page, R. H. Miller, and D. R. Keeney.) (American SOciety of Agronomy: Madison, WI.) Paton, T. R., Humphreys, G. S., and Mitchell, P. B. (1995). 'Soils: A New Global View.' (University College London Press: London.) Rayment, G. E., and Higginson, F. R. (1992). 'Australian Laboratory Handbook of Soil and Water Chemical Methods.' (Inkata Press: Melbourne.) Soil Survey Staff (1992). 'Keys to Soil Taxonomy.' 5th Edn. SMSS Technical Monograph No. 19. (Pocahontas Press: Blacksburg, VA.) Tasmania Department of Mines (1965). 'Geological Atlas 1 :63360 Series-.' Walkley, A" and Black, L A. (1934). An examination of the Degtjareif method for determining soil organic matter and a proposed modification of the chromic acid titration method~ Soil Science 31, 29-38.

Manuscript received 15 July 1996, accepted 15 August 1997 Appendix 2 Blakemore, R.J. and Kingston, T.J. (1997)

JOURNAL OF NATURAL HISTORY, 1997,31, 1683-1708 I

Opisthogastric earthworms (Megascolecidae: Oligochaeta) and allied forms in north-western Tasmania

R. J. BLAKEMORE and T. J. KINGSTON Queen Victoria Museum, Wellington St, Launceston, Tasmania, 7250

(Accepted 21 May 1997)

A new Nexogaster is established for a Tasmanian megascolecid earthworm, Nexogaster sexies sp. nov., that has an oesophageal gizzard and intestinal gizzards, the opisthogastric condition, that in this species are moniliform. The only prev­ iously known opisthogastric species from Australia, Hickmaniella opisthogaster, is also from Tasmania but has a single intestinal gizzard; this species is redescribed. A new species, Hickmaniella gogi, is bithecal but otherwise similar to the type­ species. Nexogaster differs from Hickmaniella in being lumbricine and having prostates that are racemose, rather than tubuloracemose (although both forms are an apomorphy of tubular prostates). Sympatric species are newly described that have close morphological similarities: one, Anisochaeta simpsonorum, is com­ parable with Hickmaniella, and two others, Notoscolex pilus and maestus, not only resemble one another but are also similar to Nexogaster. A major morphological difference is that M. maestus has tubular prostates; this combined with its other primitive character states, such as lumbricine setae, give an indication of a possible precursor to the other species. Extensive generic reviews and redefinitions have been required to accommodate the new species.

KEYWORDS: Anisochaeta, Cryptodrilus, Hickmaniella, Megascolex, Megascolides, Nexogaster, Notoscolex, Tasmanian earthworms.

Introduction Hickmaniella was believed by Jamieson (1974) to be unique among the known Australian Megascolecidae in possessing an intestinal gizzard, the 'opisthogastric' condition. Since 1991, a programme designed to sample native earthworms compre­ hensively throughout Tasmania for the first time has been undertaken by the second author. The material accumulated from some 600 localities has been examined by the authors and found to contain approximately 250 varieties, including several opisthogastric forms and their allies some of which are the subjects of this paper. Before these species can be described it is necessary to critically review several genera, especially those in which these species fit best. Following the taxonomic principle of priority, this review considers several 'classical' genera that have been variously synonymized. The fate of type-species is of particular concern, as the species is the basic element of morphological taxonomy.

Materials and methods Segments are counted in Arabic numerals, intersegments designated by a slash (e.g. 1/2), variations shown by a comma and range by a dash (e.g. 3,4-5). Setae,

0022-2933/97 $12-00 © 1997 Taylor & Francis Ltd. 1684 R. J. Blakemore and T. J. Kingston

counted from the ventral-most on each side, have lower case letters (a, b, c, d) and ranges are shown without a space (e.g. ab); setal lines refer to longitudinal setal series. New material is lodged in the Queen Victoria Museum, Launceston and has a QVM registration number (beginning QVM:14:), in the National Earthworm Collection of the CSIRO Division of Entomology, Canberra (prefixed ANIC:), and in the Tasmanian Museum, Hobart (TM:). The Natural History Museum, London is abbreviated BM. For material examined, H refers to holotype and P to paratype. Specimens were anaesthetized with dilute ethanol before being fixed in 10% formalin and preserved in 80% ethanol. Dissections and camera lucida drawings were by the principal author; in the figures all scale bars are 1 mm. As part of the process of determining to which genera the new species described in this paper should be allocated it became necessary to review and revise the diagnoses of six genera. For most of the individual species, allocation to genus was based upon the outcome of several generic reviews; for this reason the generic reviews are presented first followed by species' descriptions.

Systematics, generic diagnoses Hickmaniella Jamieson, 1974. Emend. Hickmaniella Jamieson, 1974: 300.

Diagnosis Setae numerous per segment. Male pores from tubuloracemose prostates paired on 18. An oesophageal gizzard in 5 and an intestinal gizzard in 19-20. Nephridia meroic (with ca. three to six tubules per side), avesiculate, not tufted anteriorly. Spermathecae one or two pairs, spermathecal diverticula clavate often with several internal chambers (i.e. multiloculate but not sessile). Calciferous glands, typhlosole and intestinal caeca absent. Penial setae present. Type-species. Hickmaniella opisthogaster Jamieson, 1974.

Distribution North/north-western Tasmania.

Remarks The generic definition is emended following re-examination of the type-species which has an intestinal gizzard in 19 and 20 (cf. 19 or 20, Jamieson) and to accommodate a new species having only one pair of spermathecae. Intestinal giz­ zards, while rare in the Megascolecidae, are found in several other families: in the Palaearctic, the Hormogastridae have oesophageal gizzards and a rudimentary intest­ inal gizzard, the Lumbricidae have only an intestinal gizzard while the Criodrilidae have oesophageal and intestinal thickenings; Neotropica1 Glossoscolecidae have a gizzard in 6 and intestinal dilation in the region of 16; the African have an oesophageal gizzard (in 5) but rarely have intestinal gizzards (Sims, 1980). These last two families have closest links with the Megascolecidae, although the intestinal gizzards may have been acquired independently (Jamieson, 1974). Hickmaniella may be derived from Anisochaeta Beddard, 1890 via its apomorphic acquisition of an intestinal gizzar:d. Especially some new species of the latter genus currently being described by the authors (see below) are superficially similar. Alternatively, it may be viewed as a derivation of a Notoscolex-like precursor that New genus Nexogaster from Tasmania 1685

has undergone transition from 8 setae per segment (the plesiomorphic lumbricine state) to more numerous setae (perichaetine state).

Anisochaeta Beddard, 1890 (Synonyms: Trichaeta, ?, ?Gemascolex) Megascolex (part) Templeton, 1844: 89; Beddard, 1890: 56; 1895: 381; Michaelsen, 1900: 212-216; 1907: 163; Stephenson, 1930: 837; Jamieson, 1974: 318-319. Perichaeta (part), Schmarda, 1861: 13; Beddard, 1890: 56. Anisochaeta Beddard, 1890: 56. Trichaeta Spencer, 1900: 30-31. Spenceriella (part? including type-species) Michaelsen, 1907: 160-161 ; Jamieson, I 972a. Gemascolex (part?) Edmonds and Jamieson, 1973: 23-24.

Diagnosis Setae> 8 per segment, at least in the hind-body. Male pores from tubuloracemose or racemose prostates paired on 18. An oesophageal gizzard in 5 (or 6). Nephridia meroic, at least in the fore-body, avesiculate, often tufted in the anterior. Spermathecae one or more pairs (sometimes unpaired?), with one or more extramural diverticula. Calciferous glands and typhlosole present or absent; intestinal caeca and gizzards absent. Penial setae present or absent. Type-species. Perichaeta coxii Fletcher, 1886: 565-659; Fletcher, 1889: 1554.

Distribution Australia (according to Beddard, 1890), New Zealand (Lee, 1959).

Remarks Australian species that would formerly have been placed in Megascolex (or in Spenceriella/Gemascolex) are allocated to the resurrected genus Anisochaeta as defined above. Beddard (1890) established Anisochaeta for three species from New South Wales, Perichaeta attenuata Fletcher, 1889, P. enormis Fletcher, 1889 and P. coxii Fletcher, 1886 (the type-species by prior description), distinguished thus: 'Setae 8 in number per segment anteriorly, afterwards increasing up to 30 [i.e. perichaetine]; nephridia diffuse [Le. meroic]; atria lobate [Le. prostates tubulora­ cemose or racemose].' This definition differed from Beddard's characterization of Megascolex principally in the transition of setae from 8 anteriorly to more numerous posteriorly. Later, Beddard (1895) transferred Anisochaeta to Megascolex where it has remained subsumed to this time. A monotypic Trichaeta was established by Spencer (1900) for Trichaeta australis from Victoria, a species that has 12 setae per segment arranged in pairs, meronephridia (tufted anteriorly) and racemose prostates. This genus was subsequently subsumed in Megascolex (e.g. by Michaelsen, 1907; Stephenson, 1930), Michaelsen (1907) defined Spenceriella with Diporochaeta notabilis Spencer, 1900 from Victoria as type-species, as he believed it differed from Megascolex principally through having plesiomorphic tubular prostates (as described by Spencer). Jamieson (1972a) (while conceding that 'the exact form [was] uncertain owing to partial fragmentation') concluded that the prostates were racemose (or 'racemose though elongate', Jamieson, 1971: 492) in a poorly preserved specimen that he designated a neotype of Diporochaeta notabilis (despite its noteworthy 1686 R. J. Blakemore and T. J. Kingston

difference in arrangement of genital markings?). This observation rendered 'Spenceriella indistinguishable from Megascolex as defined by Michaelsen (1907)'. Jamieson (l972a) omitted to define Megasco/ex but he saw no reason to consider D. notabilis to be congeneric with the type-species of Megasco/ex and therefore maintained both genera. However, if the justifiable view (e.g. held by Michaelsen, 1907: 160; Stephenson, 1930: 836) that an increase in setae in any part of the body beyond the plesiomorphic 8 per segment is the dividing line, then on the basis of its setal pattern and non-tubular prostates, D. notabi/is conforms with previous definitions, not only of Megascolex but of both Trichaeta and Anisochaeta also. Since Anisochaeta is the priority genus, Spenceriella s. Jamieson as well as Trichaeta are, necessarily, synonymized in this genus upon separation from Megascolex s. str. The type-species of Megascolex, M. caeruleus Templeton, 1844 from Sri Lanka, lacks extramural spermathecal diverticula and on this character alone . can be separated from all known Australian megascolecids. It is therefore proposed to thus restrict Megascolex and to transfer pertinent Australian species to Anisochaeta. Many Australian species that formerly would have been included in Megascolex sensu lato have been variously allocated to Spenceriella sensu Jamieson, 1972a, to Gemascolex Edmonds and Jamieson, 1973 (which differs from Spenceriel/a principally by having genital markings, where present, intersegmentally) or to Propheretima Jamieson, 1995. This last genus was stated by Jamieson (1995: 590) to differ from Spenceriella by a combination of characters: presence of setae between the male pores, restriction of the clitellum to segments 14-16 and, typically, a single female pore. However, Jamieson (l972a), in his redescription of D. notabilis has the clitellum 'poorly developed, annular, on 14-16' and in his figure (Jamieson, 1972a: fig. ID) appears to show a seta between the male pores. Moreover, at least one species included in Spenceriel/a s. Jamieson, S. conondalei Jamieson, 1995, has a single female pore, an erstwhile diagnostic characteristic of both Propheretima and Gemascolex. Full review of these genera is beyond the scope of the present study, but Anisochaeta as defined above probably forms a polyphyletic group (Stephenson, 1930: 712) and many included species may therefore eventually be transferred to other genera, including possibly several new ones.

Nexogaster gen. nov. Diagnosis Setae 8 per segment. Dorsal pores present. Male pores from racemose prostates paired on 18. An oesophageal gizzard in 5 and moniliform intestinal gizzards in the region of 22-27. Nephridia meroic, avesiculate, not tufted anteriorly. Spermathecae two pairs, sperrnathecal diverticula single, clavate. Calciferous glands and caeca absent, typhlosole present. Penial setae present. Type-species. Nexogaster sexies sp. nov. (see below)

Distribution North-western Tasmania.

Remarks Nexogaster is distinguished from all known megascolecids by the possession of an oesophageal gizzard in segment 5 and moniliform intestinal gizzards in 22,23·-26,27. The monotypic genus Pleionogaster Michaelsen, 1892 from the New genus Nexogaster from Tasmania 1687

Philippines (and doubtfully from Moluccas), while having intestinal gizzards in 26-28,29, differs principally by being perichaetine, vesiculate meroic and having an oesophageal gizzard in segment 8 (Easton, 1979). Its similarity in other respects supports a close relationship between Australian and Oriental megascolecids. Multiple intestinal gizzards are also found in the primitive and distantly-related, Oriental family Moniligastridae. Phylogenetic affinities of the present genus are unlikely to be close to those of Hickmaniella (q.v.) because taxonomic differences, such as the lumbricine setae, racemose prostates and typhlosole, suggest that their intestinal gizzards represent homoplastic rather than homologous development. Conversely, two new species (of NOlosco/ex and Megascolides) are described below which bear superficial similarity to Nexogaster sexies and may be indicative of its antecedents. For reasons that become apparent below, before Notosco/ex and Megascolides can be considered it is necessary to reaffirm Cryptodrilus.

Cryptodrihls Fletcher, 1886 (Synonym: Trinephrus) Cryptodrilus Fletcher, 1886: 570. Cryptodrilus (part) + Trinephrus; Beddard, 1895: 483. Notoscolex (part) + Trinephrus; Michaelsen, 1900: 185. Notoscolex (part); Michaelsen, 1907: 162; Stephenson, 1930: 836-837; Gates, 1959: 254; Jamieson, 1972b: 518. Cryptodrilus; Jamieson, 1972c: 154-155. Cryptodrilus (part); Jamieson, 1974: 266.

Diagnosis Setae 8 per segment. Dorsal pores present or absent. Male pores from tubulorace­ mose or racemose prostates paired on 18. An oesophageal gizzard in 5. Nephridia vesiculate meroic, with multiple bladders in at least some segments of body. Spermathecae two (or three) pairs, with one or more clavate diverticula. Extramural calciferous glands and caeca absent, typhlosole absent or present. Penial setae present or absent. Type-species. Cryptodrilus rusticus Fletcher, 1886: 570-573.

Distribution New South Wales, Victoria, Tasmania.

Remarks The type species, Cryptodrilus rusticus, has vesiculate meronephridia (i.e. several nephridia per segment with terminal bladders), tubuloracemose prostates and two (opposed) or three spermathecal diverticula. Other congeneric species have the more usual single diverticulum. The attainment of nephridial bladders serves to separate Cryptodrilus from the prior Australian genus Notosco/ex Fletcher, 1886 which in addition typically has racemose prostates and extramural calciferous glands. The genus Trinephrus (with CryptodrilusJastigatus Fletcher, 1889 as type-species) was erected by Beddard (1895) for those cryptodrilids with three pairs of nephridia per segment. Simultaneously and incorrectly Beddard placed Notosco/ex, a prior genus, as a junior synonym in Cryptodrilus. While correcting and reversing this lapse, Michaelsen (1900) included the type-species of Cryptodrilus in NOlosco/ex and placed some Tasmanian cryptodrilids, viz. Cryptodrilus officer;, c. polynephricus and Megascolides simsoni, all of Spencer (1895), in Trinephrus. Later, Michaelsen (1907) 1688 R. J. Blakemore and T. J. Kingston

transferred Trinephrus into the synonymy of Notoscolex. Whereas Jamieson (1972b: 518) included C. rusticus (but not C. fastigatus, cf. Jamieson, 1973: 235) in Notoscolex, Jamieson (l972c) restored Cryptodrilus (keeping Trinephrus as its junior synonym) for vesiculate meroic species after demonstrating multiple nephridial blad­ ders in syntypes of both C. rusticus and C. fastigatus. Subsequently, Jamieson (1974: 266) expanded the diagnosis of Cryptodrilus, ignoring variations in nephridia from vesiculate to avesiculate (and presence or absence of preseptal funnels) and ignoring prostate form from tubular to racemose, so that it subsumed those characters that define both Notoscolex and Megascolides. This action is here reversed as conformity to types requires that species with plesiomorphic tubular prostates, never part of the original descriptions of Cryptodrilus, are excluded and only vesiculate meroic species are allowed. Cryptodrilus is diagnosed above and Notoscolex and Megascolides are restored and redefined below.

Notoscolex Fletcher, 1886 (Synonyms: Tokea, Pseudonotoscolex, ?Oreoscolex). NOloscolex Fletcher, 1886: 546. NOloscolex + Megascolides (part); Michaelsen, 1900: 187. Notoscolex (part); Stephenson, 1930: 836. Notoscolex; Michaelsen, 1907: 160-162; Lee, 1959: 317; Gates, 1959: 254; Jamieson, 1973: 236,239. Tokea Benham, 1904: 240. Pseudonotoscolex Jamieson, 1971: 496. Oreoscolex (part? excluding type-species); Jamieson, 1973: 215-252; 1974: 302-303.

Diagnosis Setae 8 per segment. Dorsal pores present or absent. Male pores from racemose (or tubuloracemose?) prostates paired on 18. An oesophageal gizzard in 5 or 6. Nephridia meroic, at least in the fore-body, avesiculate, sometimes tufted. Spermathecae two or three pairs (or unpaired?), with multiloculate sessile (or one or more clavate) diverticula. Typhlosole typically absent; extramural calciferous glands typically present; intestinal caeca absent. Penial setae typically present. Type-species. No toscolex camdenensis Fletcher, 1886: 546-551, PI. VII, figs 1-5.

Distribution New South Wales, Victoria, Tasmania, , southern Queensland, (New Zealand, ?South India, Sri Lanka).

Remarks For Notoscolex, the 'classical' definition of Michaelsen (1900, 1907) and Stephenson (1930) (as perpetuated by Lee, 1959 and Gates, 1960) is retained for Australian species, after removal ofvesiculate species to Cryptodrilus (q.v.). Although Jamieson (1973: 239) suggested that Notoscolex should be restricted to species with extramural calciferous glands in 14-16 (?and/or 9-13 as in N. montiskosciuskoi Jamieson, 1973), no significance was attached to the fact that the type-species has multiloculate spermathecal diverticula, often a generic character. It is possible that only those species with multioculate diverticula truly belong in Notoscolex. In 1973, Jamieson erected the monotypic genus Oreoscolex for his Oreoscolex imparicystis, a species lacking calciferous glands but possessing these diagnostic features: lumbricine setae, avesiculate meronephridia, racemose prostates and New genus Nexogaster from Tasmania 1689 unpaired, bidiverticulate spermathecae. An expanded diagnosis -of Oreoscolex by Jamieson (1974: 302) permitted presence or absence of calciferous glands, racemose or tubuloracemose prostates, and uniloculate or multiloculate spermathecal divertic­ ula. This re-definition was so expansive that it allowed synonymy with Notoscolex, a prior genus (and also with Pseudonotoscolex?). This impacts especially upon two of the eight Tasmanian species that Jamieson (1974) included under Oreoscolex: Notoscolex leaf Michaelsen, 1910 and N. sexthecatus (Jamieson, 1974), as well as Cryptodrilus ? albertisii (Cognetti, 1910), Notoscolex officeri (Spencer, 1895) and N. simsoni (Spencer, 1895), as the states of the caudal nephridia in these species ­ pivotal for designation in Jamieson's scheme - were inconclusive. Oreoscolex is herein considered a junior synonym of Notoscolex, although an alternative view could allow its retention as a monotypic genus holding only the type-species, 0. imparicystis, with its unpaired spermathecal pores. Two Notoscolex species have recently been described by Jamieson (1995), one as Cryptodrilus bunyaensis from southern Queensland, the other as Oreoscolex retrocystis (represented by a posterior amputee, acIitellate holotype that is possibly the 'Notoscolex sp. from Galston Gorge' as adjudged in Jamieson, 1972c). Jamieson (1971) erected the monotypic Pseudonotoscolex (a genus similar to Notoscolex) for his P. pallinupensis from Western Australia, a species represented by a posterior-amputee holotype. Later, Jamieson (1981: 907) raised the possibility of absorbing Pseudonotoscolex in Cryptodrilus, a move not permissible for an avesic­ ulate species although its characteristics comply with the diagnosis of Notoscolex above. Pseudonotoscolex is therefore synonymized under Notoscolex. Benham's New Zealand genus Tokea was synonymized under Notoscolex in Stephenson (1930: 837) (cf. Lee, 1959: 259), but it appears that some included species having tubular prostates actually belong in Megascolides.

Megascolides McCoy, 1878. Emend (Synonyms: Austrohoplochaetella, Pseudocryptodrilus) Megascolides McCoy, 1878: 21. Cryptodri/us (part); Beddard, 1895: 497. Megascolides; Michaelsen, 1900: 182; 1907: 160-161. Megascolides + Notoscolex (part); Stephenson, 1930: 835. Megasco/ides; Lee, 1959: 284-285. Austrohoplochaetella Jamieson, 1971: 490. Pseudocryptodrilus Jamieson, 1972c: 172.

Diagnosis Setae 8 per segment. CIitelIum typically developed over at least four segments. Male pores from tubular prostates paired on 18. An oesophageal gizzard in 5 or 6 (or 7). Nephridia meroic, at least in the fore-body, avesiculate or vesiculate, some­ times tufted anteriorly. Spermathecae one or more pairs, each with a clavate divertic­ ulum. Typhlosole and calciferous glands present or absent; intestinal caeca absent. Penial setae present or absent. Type-species. Megascolides australis McCoy, 1878.

Distribution Victoria, New South Wales, Tasmania, Western Australia, (New Zealand, ?North America). 1690 R. J. Blakemore and T. J. Kingston

Remarks The Victorian type-species, Megascolides australis, has numerous meronephridia per segment whereas other species included in the genus have fewer tubules and some appear transitional from holonephridia to meronephridia. In the present scheme, the rationale is that any nephridial forms intermediate between holonephri­ dia and meronephridia (with the exception of tufted pharyngeal holonephridia) are classed as apomorphic meroic derivations. This pragmatic approach is consistent with Michaelsen and with Stephenson (1930: 830) who stated: 'From Plutellus is derived Megascolides, in which the nephridia are breaking up or have broken up; this apparently does not always take place in the same way. In one group of forms there are three or four nephridia on each side of each segment, all about the same size, while in other cases there is one large one and a number of quite small ones; however, all stages of the process are included in this genus, so-long as the prostates and setae retain their original condition.' The generic definition above is emended only to note that both vesiculate and avesiculate nephridial forms are included since Jamieson (19nc; 1973) found multiple bladders in Megascolides tenuis (Fletcher, 1889). Incidentally, Michaelsen (1907: 161) had already transferred this New South Wales species to this genus (cf. Jamieson). Megascolides is distinguished from Notoscolex and Cryptodrilus by its possession of plesiomorphic tubular prostates. Jamieson (1971) erected the monotypic genus Austrohoplochaetella for his species A. kendricki from Western Australia, its diagnosis differing from that of Megascolides principally because it exhibited transition from meronephridia to 'a pair of single stomate megameronephridia' caudally. Jamieson (I9nc) proposed the monotypic genus Pseudocryptodrilus for Megascolides diaphanus Spencer, 1900 from Victoria which has a similar transition, but this time from meronephridia to 'a pair of stomate exonephric holonephridia' caudally. Both of these nephridial states, if indeed they differ, were preallowed in Megascolides by Michaelsen (1907: 160, 'wenigstens im Vorderkorper mikronephridisch'-at least in the fore-body meroic), as he described Megascolides nokanenaensis from Western Australia having just such nephridial transition. This is perhaps why Jamieson (1981: 907) stated that Austrohoplochaetella is probably a junior synonym of Megascolides, a view supported here. If this is the case, there is little justification for retaining Pseudocryptodrilus and it is therefore similarly synonymized. Two Tasmanian species were placed in Megascolides by Spencer (1895), but neither belonged as Notoscolex simsoni (Spencer, 1895) has non-tubular prostates and Vesiculodrilus? bassanus (Spencer, 1895) was described with (vesiculate?) holonephridia.

Systematics, species descriptions

Hickmaniella gog; sp. nov. (Fig. 1) Material examined HOLOTYPE (H) QVM:14:3328, Gog Range, N. Tasmania, 41°29'S 146°27'E, 27 August 1991, T. J. Kingston (mature, dissected and figured). PARATYPES (PI) ANIC:RB.96.11.l, same details as H (mature, dissected); (P2) ANIC:RB.96.Il.1, same details as H (immature); (P3) QVM:14:3319, same details as H (immature, dissected). New genus Nexogaster from Tasmania 1691

(c)

(d)

FIG. l. Hickmaniella gogi sp. nov., holotype: (a) prostate and penial setae; (b) spermatheca; (c) prostomium; (d) intestinal gizzards in 19-20(t21).

(P4) TM:KI523, Little Fisher River, Lake Rowallan, N.W. Tasmania, 41042'S 146°18'E, 6 October 1992, R. D. D'Orazio and M. Cooper (mature, dissected and figured); (P5·-9) QVM:14:0416, details same as P4 (five matures, one aclitellate). (PI0-14) QVM:14:0417, Dublin Creek, Lake Rowa11an, N.W. Tasmania, 41°42'S 146°15'E, 6 October, 1992, R. D. D'Orazio and M. Cooper (five specimens: two clitellate, one ac1itellate, two immature). 1692 R. J. Blakemore and T. J. Kingston

(PI5) QVM:14:3532, Pelion Gap, N.W. Tasmania, 41°52'S 146°03'E, 31 January, 1992, D. Baker (mature, dissected). (PI6-18) ANIC:RB.96.11.3, Snake Creek Road, Mole Creek, N.W. Tasmania, 41°37'S 146°16'E, 5 October, 1992, R. D. D'Orazio and M. Cooper (one mature and two sub-adults, one dissected); (PI9-21) QVM:14:0970, same details P16-18 (one mature and two sub-adults). (P22-23) TM:Kl 524-1 525, O'Neils Road, Gowrie Park, N.W. Tasmania, 41°26'S 146°14'E, 23 November 1992, R. D. D'Orazio and M. Gittus (one mature and one sub-adult, dissected); (P24-25) QVM:14:0679, same details P22-23 (two sub-adults).

External features Body stout hardly tapering to tail, first segment compressed. Length mm: (H) 75, (PI) 70, (P4) 60. Width: ca 7 mm. Segments: (H) 98, (PI) 97, (P2) 91, (P3) 70, (P4) 103. Colour: unpigmented in alcohol, c1itellum buff. Prostomium: tapering epilobous, faintly furrowed dorsally. Clitellum: weak in t 13,14-17 (H, PI) or tumid in tI3,14-17,t18 (most other matures). Dorsal pores: rudimentary in 2/3, open from 3/4. Nephropores: not found. Setae: numerous, 40~48 per segment, ventral couples ab in regular rows, other rows sinuous; dorsal setal gap ca !aa. Spermathecal pores: paired in 8/9 in a or ab lines within tumid area. Female pores: widely paired on 14. Male pores: paired on 18 on flat, irregular pads with two or three penial setae protruding. Genital markings: median troughs in bb in 10/11 (PIO-I2) or 13/14 and/or 16/17 (PI, P4-P9, PI6-22); two pairs of ill-defined hollow disks joined by median troughs in 17/18 and 18/19 wider apart than the male pores (H, PI and all other matures); elongate, depressed pads in bb in 19/20 same width as male pores (H, PI and all matures except PI0-15, P22) ..

Internal anatomy Septa: 7/8-12/13 thickened, 9/10/11/12 peripherally adpressed in H (as pericar­ diac testis-sacs?). Gizzards: muscular but compressible oesophageal gizzard in 5; large muscular intestinal gizzard in 19-20 and perhaps part of 21 also modified; externally with lateral bands of muscle fibres, internally with longitudinal striations, wall ca 0'5mm thick (thicker than that of oesophageal gizzard). Oesophagus: not especially dilated. Nephridia: avesiculate meroic, five or six tubules per side equatori­ ally connected by mesentery; larger anteriorly but not tufted; funnels not found. Vascularization: dorsal blood vessel single; hearts 10-12 with connectives to supra­ oesophageal vessel in 7-14. Spermathecae: one pair in 9; ampulla saccular (smooth in H, PI; irregular in outline in some other matures) on thick duct bearing small clavate diverticulum with numerous iridescent chambers internally. Male organs: holandric, iridescent testes in mucus in 10 and 11; small racemose seminal vesicles in 9 and 12. Ovaries: compact in 13. Prostates: tubuloracemose in 18, gland folded over itself; overlain by long penial setal sheaths and tendons. Intestine: from 18, dilated and thin-walled on either side of gizzard in 18 and in 21, spiralling from 22; typhlosole absent; gut contains soil with numerous quartz grits and sand.

Remarks The definition of Hickmaniella has been emended to accommodate this bithecate species. Specific differences that further separate Hickmaniella gogi from H. opisthogaster are position of first dorsal pore and the form and arrangement of New genus Nexogaster from Tasmania 1693 the genital markings. The genus Hickmaniella is perichaetine with tubuloracemose prostates, an intestinal gizzard in 19-20 and lacks a typhlosole (on these characters it differs from Nexogaster gen. nov. described below).

Distribution and habitat Gog Range, Lake Rowallan, Pelion Gap, Mt. Roland and Mole Creek in north/ north-western Tasmania.

Hickmaniella opisthogaster Jamieson, 1974 (Fig. 2) Hickmaniella opisthogaster Jamieson, 1974: 301-302, Fig.18A (p.270), 32C, D (p.325), PI. 64-66.

Material examined HOLOTYPE: (H) TM:K360, Parrawe, 41°18'S 145°35'E, 25 August, 1954, J. L. Hickman (mature, dissected). PARATYPES: (PI) BM:1973.2.34, same details as H (mature, dissected); (P2) TM:K361, same details as H (immature, here dissected to confirm intestinal gizzard). Specimens. QVM:14:1619, Stephens Rivulet, Balfour Track Forest Reserve, N.W. Tasmania, 41° 08'S 144° 57'E, 18 May 1993, R. D. D'Orazio and D. E. Soccol (mature, dissected and figured, plus an immature). QVM:14:l616, Bond Tier South, Dismal Swamp, N.W. Tasmania, 40 0 58'S 144°51 'E, 19 May 1993, R. D. D'Orazio and D. E. Soccol (five specimens, one immature); ANIC:RB.96.1 1.4, same details, (six matures, one dissected); QVM:14:3329-3330, Dismal Swamp, N.W. Tasmania, 40 0 59'S 144°51'E, 8 September 1987, T. J. Kingston (two matures, one dissected). QVM:14: 1260, Tram Road Picnic Area, Calder, N.W. Tasmania, 4P02'S 145°41'E, 19 April 1993, R. D. D'Orazio and D. E. Soccol (seven specimens). QVM:14:1080, Hellyer Gorge Reserve, N.W. Tasmania, 41°17'S 145°37'E, 31 May 1993, R. D. D'Orazio and D. E. Soccol (one mature and one immature). QVM:14:1074; Belmont Road, Waratah, N.W. Tasmania, 41023'S 145°32'E, 31 May 1993, R. D. D'Orazio and D. E. Soccol (three matures). QVM:14:562, West Calder Road, Calder, N.W. Tasmania, 41°05'8 145°37'E, 19 April 1993, R. D. D'Orazio and D. E. Soccol (one specimen). QVM:14:1613, Trowutta Caves State Reserve, Smithton, N.W. Tasmania, 41°04'S 145°06'E, 17 May 1993, R. D. D'Orazio and D. E. Soccol (one mature, dissected). QVM:14:3277, Walking Track off Bass Highway, near Dismal Swamp, 40 0 57'S 144°49'E, 24 June 1993, T. J. Kingston, J. Buckerfield, R. J. Blakemore (two matures); ANIC:RB.96.11.5, same details (two matures, one dissected). QVM:14:3554, Fern Glade Reserve, Burnie, N.W. Tasmania, 41°05'S 145°55'E, 3 December 1996, R. J. Blakemore and T. J. Kingston (three matures, one dissected). Type material not examined (P3--4) AM: W5322-5323, Table Cape, 400 57'S 145°43'E, 24 August 1954, J. L. Hickman.

1?xternalJfeatures Body short and robust, first segment compressed. Lengths mm: 45-70. Width: ca 5-5'5 mm. Segments: 80--90. Colour: unpigmented, clitellum buff. Prostomium: epilobous, weakly furrowed to appear tanylobous. Clitellum: t 13-!17 (often appears saddle-shaped when markings impinge). Dorsal pores: from 4/5 (rudimentary in 3/4 1694 R. J. Blakemore and T. J. Kingston

FIG. 2. Hickmaniella opisthogaster, specimen QVM:14:1619: (a) prostate and penial setae; (b) sperrnathecae; (c) prostomium; (d) intestinal gizzard in 19-20.

in some specimens). Nephropores: not found. Setae: 40-50 per segment; ventral gap wide, dorsal gap less so. Spermathecal pores: in 7/8/9 in a or ab lines. Female pores: paired on 14. Male pores: paired on 18 on prominent mounds, just wider than setal gap. Genital markings: large, paired (or analogue) glandular mounds in 11/12 and, New genus Nexogaster from Tasmania 1695 variously, in 10/11, 14/15-16/17 and 19/20-22/23, slightly wider than male and spermathecal pores.

Internal anatomy Septa: 6/7--12/13 thickening. Ventral nerve cord large. Gizzards: large spherical but weakly muscular oesophageal gizzard in 5; intestinal gizzard large and muscular in 19-20. Oesophagus: not especially dilated. Nephridia: avesiculate meroic, three or four per side equatorial1y as small, convoluted masses; not tufted anteriorly; funnels not found. Vascularization: hearts 10-12; supra-oesophageal vessel moder­ ately developed in 7-13, large in 14-17. Spermathecae: two pairs in 8 and 9; saccular ampulla on short duct, diverticulum clavate with numerous internal chambers (some­ times rosette-like but not sessile). Male organs: holandric, testes and funnels in 10 and II, iridescent and invested in mucus; racemose seminal vesicles in 9 and 12. Ovaries: in 13. Prostates: tubuloracemose, folded once in IS; long penial setae present. Intestine: from IS, dilated and thin-walled atria on either side of gizzard in IS and 21 (the anterior crop-like dilation often impinges on gizzard in 19; that in 21 sometimes extends into segment 22 also), intestine spirals from 22; typhlosole absent; gut contains soil.

Remarks This account confirms and considerably augments the type description. Morphological features recorded by the current study are the compressed first segment, epilobous prostomium, lack of pigmentation, lesser extent of the clitellum, variations in the distribution of genital markings, the large ventral nerve cord and the intestinal gizzard in 19 and 20 (rather than 19 or 20 according to Jamieson, 1974) with intestinal dilations on either side in 18 and 21. The distributional range is also expanded.

Distribution and habitat North-western Tasmania from a north-south line from Bumie-Waratah, west­ wards to Balfour and Marrawah, including Parrawe, Hellyer Gorge, Wynyard, Smithton. Found mainly in wet sclerophyll and rainforest soils.

Anisochaeta simpsonorum sp. nov. (Figs 3,4)

Material examined HOLOTYPE: (H) QVM:14:3275, Dismal Swamp Nature Reserve, N.W. Tasmania, 40 0 59'S 144°51'E, 8 September 1987, T. J. Kingston (mature, posterior amputee, dissected and figured). PARATYFES: (PI) ANIC:RB.96.11.6, Walking Track off Bass Highway, near Dismal Swamp, 40 0 57'S 144°49'E, 24 June 1993, T. J. Kingston, J. Buckerfield, R. J. Blakemore (mature, posterior amputee, dissected and figured). (P2-4) TM:K1526-152S, Belmont Rd., Waratah, N.W. Tasmania, 4I023'S 145°32'E, 31 May 1993, R. D. D'Orazio and D. E. Socco! (3 matures, P2 dissected and figured); (P5-PS) ANIC:RB.96.11.7, same details (4 matures); (P9-11) QVM:14:I075, same details, (three matures, PI 1 dissected, plus three unregistered juveniles). 1696 R. 1. Blakemore and T. 1. Kingston

(c)

FIG. 3. Anisochaeta simpsonorum sp. nov., holotype: (a) racemose prostate; (b) spermathecae; (c) prostomium. New genus Nexogaster from Tasmania 1697

FIG. 4. Anisochaeta simpsonorum sp. nov., variations in spermathecae and markings: (a) PI; (b) P2.

(P12) QVM:14:3529, 'Killara' property, Marrawah, N.W. Tasmania, 41°01 'S 144.o44'E, 7 September 1987, T. J. Kingston (mature). (P13) QVM:14:3530, Wombat Hill near Waratah, N.W. Tasmania, 4lo29'S 145°27'E, 22 September 1990, R. Mesibov (mature, posterior amputee, dissected). (P14-16) ANIC:RB.96.12.3-5, 1.5 Km along forest track in Crown land east of 'Killara' property, Marrawah, 4lo01'S 144°44'E, 4 December 1996, R. J. Blakemore and T. J. Kingston (three matures, one dissected); (PI7-l9) TM:K1535-l537, same details, (three matures, one dissected); (P20) QVM:14:3572, same details (one mature, dissected; sample also contains nine sub-adult and immature specimens that superficially agree). (P21) QVM:14:3052, Corinna Rd., N.W. Tasmania, 41°36'S l45°08'E, 2 June 1993, R. D. D'Orazio and D. E. Soccol (mature, dissected-separated off 14:1052 which contained 18 specimens). (P22) QVM:14:1048, Pieman River, Corinna, N.W. Tasmania, 41°39'S 145°05'E, 2 June 1993, R. D. D'Orazio and D. E. Soccol (mature, dissected).

External features Lengths mm: range 60-115; (P2-3) 93, (P5-12) 80--90, (PI 4-20) 60-80, (P21-22) 90-115, (H and PI are 50 + and 55 + mm, respectively). Segments: (P2) 136, (P3) 140 (P2l) 124, (P22) 114. Colour: pale, unpigmented or with a yellow 1698 R. J. Blakemore and T. J. Kingston

tinge, clitellum orange/buff. Prostomium: open epilobous. Clitellum: .t13-17. Dorsal pores: from 4/5. Setae: 12 throughout in mostly regular rows except for some irregularities in cd lines posteriorly. Nephropores: not seen. Spermathecal pores: paired in 7/8-8/9 near a setal lines. Female pores: paired anteriomedial to setae a. Male pores: paired near a setal lines on small raised papillae within rimmed, mid­ ventral tumid area. Genital markings: paired pads centred in b lines and extending slightly beyond ab lines in 11/12; (12/13 infolded ventrally in several specimens); smaller pads paired (or analogue) in ab lines in 16/17, sometimes absent; elongate trough in bb or aa in 19/20 surrounded by tumid area extending to setal arcs of 19-20; paired or single darkened patches in aa in 20/21. Markings are consistently in 11/12 and 19/20, however, PlO-13 and P21-22 have aberrant markings: in PlO and P12 they are asymmetrical (in some of 11/12/13,16/17 and 18/19/20/21), while Pll, P13, P21-22 have elongate troughs in bb in 11/12 (and in aa in 16/17 in PIl, P21-22) and in aa in 19/20 (Pl1, P13, P21-22).

Internal anatomy Septa: thin in anterior, 9/10--12/13,13/14 only slightly thickened. Gizzard: in 5, muscular and barrel-shaped with anterior flange, displaced posteriorly. Oesophagus: dilated and internally lamellate in 10-16, especially 15-16, but not calciferous. Nephridia: avesiculate meroic, ca two or three small clusters of tubules per side becoming laterally connected bands in midbody; not tufted anteriorly; funnels not found. Vascularization: dorsal blood vessel single; hearts in 10-12; supra-oesophageal vessel in 9-13. Spermathecae: two pairs in 8 and 9; ampulla saccular on longer duct with clavate diverticulum from middle of duct (iridescent bulb sometimes has irregular outline). Male organs: holandric, iridescent testes and funnels in mucus in 10 and 11; seminal vesicles paired (sometimes rudimentary), saccular in 9 and racemose in 12. Ovaries: large in 13; small ovisacs paired in 14. Prostates: in 18, racemose, bilobed but appear as rosettes; ducts sinuous, wide at base; penial setae not found. Intestine: origin 18 (appears to be 17 in some specimens where septum 17/18 deflected anteriorly); darker and more dilated from 19; deep typhlosole developing from 20; intestinal gizzards absent; gut contains soil, some organic matter and quartz grits.

Remarks This species is morphologically similar to parts of an Anisochaeta montisarthuri (Jamieson, 1974) species-complex at present under investigation in this laboratory. Differences include the numbers of setae, the positions of the genital markings, spermathecal and male pores, and the presence of seminal vesicles in 9.

Distribution and habitat North-western Tasmania, in rainforest or wet sc1erophyll forest soils. Samples from the Bass Highway, Belmont Rd and Dismal Swamp sites also contained specimens of Hickmaniella opisthogaster, a species having similar genital markings in 11/12.

Nexogaster sexies gen. et sp. nov. (Fig. 5) l\tlaterial examined HOLOTYPE (H) QVM:14:3320, 'Killara' property, Marrawah, N.W. Tasmania, 41°01'S 144°44'E, 7 September 1987, T. J. Kingston (mature, dissected and figured). New. genus Nexogaster from Tasmania 1697

FIG. 4. Anisochaeta simpsonorum sp. nov., variations in spennathecae and markings: (a) PI; (b) P2.

(P12) QVM:14:3529, 'Killara' property, Marrawah, N.W. Tasmania, 41 0 01'S 144°44'E, 7 September 1987, T. J. Kingston (mature). (P13) QVM:14:3530, Wombat Hill near Waratah, N.W. Tasmania, 41 0 29'S 145°27'E, 22 September 1990, R. Mesibov (mature, posterior amputee, dissected). (P14-16) ANIC:RB.96.12.3-5, 1.5 KIn along forest track in Crown land east of 'Killara' property, Marrawah, 41 °Ol'S 144°44'E, 4 December 1996, R. J. Blakemore and T. J. Kingston (three matures, one dissected); (P17-19) TM:KI535-1537, same details, (three matures, one dissected); (P20) QVM:14:3572, same details (one mature, dissected; sample also contains nine sub-adult and immature specimens that superficially agree). (P21) QVM:14:3052, Corinna Rd., N.W. Tasmania, 41 0 36'S 145°08'E, 2 June 1993, R. D. D'Orazio and D. E. Soccol (mature, dissected-separated off 14:1052 which contained 18 specimens). (P22) QVM:14:1048, Pieman River, Corinna, N.W. Tasmania, 41 0 39'S 145 Q 05'E, 2 June 1993, R. D. D'Orazio and D. E. Soccol (mature, dissected).

External features Lengths mm: range 60-115; (P2-3) 93, (P5-12) 80-90, (PI4-20) 60-80, (P21-22) 90-115, (H and PI are 50 + and 55 + mm, respectively). Segments: (P2) 136, (P3) 140 (P21) 124, (P22) 114. Colour: pale, unpigmented or with a yellow 1698 R. J. Blakemore and T. 1. Kingston tinge, clitellum orange/buff. Prostomium: open epi10bous. Clitellum: 113-17. Dorsal pores: from 4/5. Setae: 12 throughout in mostly regular rows except for some irregularities in cd lines posteriorly. Nephropores: not seen. Spermathecal pores: paired in 7/8-8/9 near a setal lines. Female pores: paired anteriomedial to setae a. Male pores: paired near a setal lines on small raised papillae within rimmed, mid­ ventral tumid area. Genital markings: paired pads centred in b lines and extending slightly beyond ab lines in 11/12; (12/13 infolded ventrally in several specimens); smaller pads paired (or analogue) in ab lines in 16/17, sometimes absent; elongate trough in bb or aa in 19/20 surrounded by tumid area extending to setal arcs of 19-20; paired or single darkened patches in aa in 20/2l. Markings are consistently in 11/12 and 19/20, however, PIO-13 and P21-22 have aberrant markings: in PI0 and P12 they are asymmetrical (in some of 11/12/13, 16/17 and 18/19/20/21), while PlI, P13, P21-22 have elongate troughs in bb in 11/12 (and in aa in 16/17 in PlI, P21-22) and in aa in 19/20 (Pll, P13, P21-22).

Internal anatomy Septa: thin in anterior, 9/10-12/13,13/14 only slightly thickened. Gizzard: in 5, muscular and barrel-shaped with anterior flange, displaced posteriorly. Oesophagus: dilated and internally lamellate in 10-16, especially 15-16, but not calciferous. Nephridia: avesiculate meroic, ca two or three small clusters of tubules per side becoming laterally connected bands in midbody; not tufted anteriorly; funnels not found. Vascularization: dorsal blood vessel single; hearts in 10-12; supra-oesophageal vessel in 9-13. Spermathecae: two pairs in 8 and 9; ampulla saccular on longer duct with clavate diverticulum from middle of duct (iridescent bulb sometimes has irregular outline). Male organs: holandric, iridescent testes and funnels in mucus in 10 and 11; seminal vesicles paired (sometimes rudimentary), saccular in 9 and racemose in 12. Ovaries: large in 13; small ovisacs paired in 14. Prostates: in 18, racemose, bilobed but appear as rosettes; ducts sinuous, wide at base; penial setae not found. Intestine: origin 18 (appears to be 17 in some specimens where septum 17/18 deflected anteriorly); darker and more dilated from 19; deep typhlosole developing from 20; intestinal gizzards absent; gut contains soil, some organic matter and quartz grits.

Remarks This species is morphologically similar to parts of an Anisochaeta montisarthuri (Jamieson, 1974) species-complex at present under investigation in this laboratory. Differences include the numbers of setae, the positions of the genital markings, spermathecal and male pores, and the presence of seminal vesicles in 9.

Distribution and habitat North-western Tasmania, in rainforest or wet sclerophyll forest soils. Samples from the Bass Highway, Belmont Rd and Dismal Swamp sites also contained specimens of Hickmaniella opisthogaster, a species having similar genital markings in 11/12.

Nexogaster sexies gen. et sp. nov. (Fig. 5)

Material examined HOLOTYPE (H) QVM:14:3320, 'IGllara' property, Marrawah, N.W. Tasmania, 41°01'S 144°44'E, 7 September 1987, T. J. IGngston (mature, dissected and figured). New genus Nexogaster from Tasmania 1699

(d)

......

FIG. 5. Nexogaster sexies sp. nov., holotype: (a) racemose prostate and penial setae; (b) spermathecae; (c) prostOlnium; (d) intestinal gizzards in 22~27. 1700 R. J. Blakemore and T. J. Kingston

PARATYPES: (PI) ANIC:RB.96.I1.9, same details as H (mature, dissected); (P2) TM:K1529, same details as H (mature); (P3) ANIC:RB.96.11.10, same details as H (mature, aclitellate); (P4), QVM:I4:3324, same details as H (a mature specimen, dissected) . (P5) QVM:14:3571, New Paddock at 'Killara', 24 June 1993, T. J. Kingston, R. J. Blakemore, J. Buckerfield, S. Pilkington (a mature specimen); (P6) ANIC:RB.96.l2.2, same details as P5 (mature, dissected); (P7), TM:K1534, same details as P5 (mature).

External features Body slender, often dorso-ventrally flattened, first segment not reduced. Length mm: range 60-90; (P6-7) 90, (H, P2 and P5) 80, (PI) 75, (P3) 65, (P4) 60. Width: ca 3·3 mm. Segments: (H) 132, (PI) 128. Colour: unpigmented in alcohol, small yellow dots laterally near c setae; clitellum buff. Prostomium: open epilobous. Clitellum:!13-16,t17. Dorsal pores: small in 4/5, larger from 5/6. Nephropores: not found. Setae: eight in regular rows (except in some specimens setae are slightly irregular at the caudal extremity and, seen in H, occasional odd or duplicated seta added); PI has several dark speckles of unknown origin ventrally on 18 and 19. Spermathecal pores: 7/8--8/9 in a lines. Female pores: paired. Male pores: paired on small mounds in abo Genital markings: large, paired sucker-like discs in 10/11 and sometimes also in 11/12 (in H, P3, P6, P7) centred in a lines; similar disks in 17/18 and some of 18/19-20/21 in ab (markings absent from 18/19 in PI, P4 and P7; absent from 20/21 in P2, P5 and P7); i.e., consistently occurring only in 10/11, 17/18 and 19/20.

Internal anatomy Gizzards: muscular oesophageal gizzard in 5; intestinal gizzards smooth, muscu­ lar and moniliform in 22-27 (in H and PI) or 23-27 (P4) or 22-26 (P6), Le. five or six of. Oesophagus: slightly dilated but not calciferous in 14---15,16. Nephridia: avesiculate meroic, clusters of four or five tubules per side centred around bc and easily broken up on dissection; larger in clitellar region, becoming smaller after this; not tufted anteriorly; funnels not found. Vascularization: hearts 10-12; supra­ oesophageal vessel 9-12. Spermathecae: two pairs in 8 and 9; saccular ampulla with large, wholly iridescent, clavate diverticulum on duct ectally. Male organs: holandric, iridescent testes in 10 and 11; racemose seminal vesicles paired in 9 and 12. Ovaries: large in 13. Prostates: racemose in 17-19,20; long, curving penial setae present. Intestine: from 18, dilated and thin-walled in 18-21,22; intestinal gizzards 22,23-26,27, moderately deep typhlosole after intestinal gizzards from 28; gut contains organic matter or dark soil and many quartz grits.

Remarks Hickmaniella (q.v.) was the only previously known Australian megascolecid to have an oesophageal and an intestinal gizzard. From this genus Nexogaster differs by being lumbricine and by having racemose rather than tubuloracemose prostates, clavate spermathecal diverticula without multiple internal chambers, a typhiosole, and 5-6 intestinal gizzards in series rather than a single intestinal gizzard. Despite literally one or two aberrant setae occurring on the tail in the holotype (at least?), these are not sustained and consequently the lumbricine state is retained. New genus Nexogaster from Tasmania 1701

The name Nexogaster sexies alludes to the connected intestinal gizzards repeated about 6 times.

Distribution and habitat Marrawah, on a farm property. Specimens were collected following cultivation of indigenous swampy heath vegetation and after sowing to pasture. This narrowly distributed species may thus be endangered by agriculture.

Notoscolex pilus sp. nov. (Figs 6, 7)

Material examined HOLOTYPE: (H) QVM:14:3325, Dismal Swamp Nature Reserve, N.W. Tasmania, 40 0 59'S 144°51'E, 8 September 1987, T. 1. Kingston (mature, dissected and figured). PARATYPES: (PI) ANIC:RB.96.11.8, same collection data as H (mature, dissected) . (P2) TM:KI530, Belmont Rd, Waratah, N.W. Tasmania, 41 '23'S 145°32'E, 31 May 1993, R. D. D'Orazio and D. E. Soccol (mature, dissected); (P3) QVM:14:3326, same details (mature); (P4) QVM:14:3566, same details (weakly clitellate mature, possibly a posterior regenerate, dissected; plus an unregistered immature that superficially agrees). (P5) QVM:14:2524, Wombat Hill near Waratah, N.W. Tasmania, 41°29'S 145°27'E, 22 September 1990, R. Mesibov (mature, dissected; (P6) ANIC:RB.96.11.14, same details (mature); (P7) ANIC:RB.96.1U5, same details (mature, dissected); (P8) TM:KI533, same details (mature); (P9), same details (mature, dissected); (PlO), QVM:14:3567, same details (mature); (PU), QVM:14:3568, same details (sub-adult); (PI2) QVM:14:3569, same details (sub­ adult). (P13), QVM: 14:0111, Frog Flats, Pelion Valley, N.W. Tasmania, 41°48'S 146°02'E, 13 February 1992, T. 1. and M. E. Kingston (mature, dissected and figured). (PI4) QVM:14:0124, near Old Pelion Hut, Pelion Valley, N.W. Tasmania, 41 °48'S l46°03'E, 12 February 1992, T. J. and M. E. Kingston (mature).

External features Lengths mm: range 30-55; CH, PI, P13-14) 30, (P2-3) 55, (P5-1O) 38-45. Width: 1-8 mm. Segments: (H and P13) 84, (PI) 95, (P2) 98, (P5) 94. Colour: unpigmented or with yellow tinge in alcohol; clitellum yellowish. Prostomium: open epilobous (faint ventral cleft on peristomium in some specimens). Clitellum: -113-16,t17. Dorsal pores: small in 4/5, larger from 5/6. Nephropores: not found. Setae: eight throughout in regular series (sometimes slightly irregular in tail). Spermathecal pores: 7/8-8/9 in a lines. Female pores: paired. Male pores: paired on small mounds in ab; upward curving tips of long penial setae protrude. Genital markings: faint paired discs posterior to spermathecal pores in 8 and 9 (H, P5-12­ these Wombat Hill specimens, except P6, PlO-l1, have additional markings in either 7 or 10); large mid-ventral pad with median eye-like marking in 10 (P4) or 13 (P5, P8-9, P12) or in 17 (H, PI, P12) or paired in 17/18 in b lines (P2-P12) or a lines (P13-14 from Pelion Valley); some specimens possibly have weak paired markings in b lines in 18/19; paired eye-like markings within tumid pads in ab in 19/20 (P3, 1702 R. J. Blakemore and T. J. Kingston

(c)

,-,---- d a h C-'

(a)

FIG. 6. Notoscolex pilus sp. nov., holotype: (a) tubuloracemose prostate with external duct and penial setae; (b) spermathecae; (c) prostoInium. New genus Nexogaster from Tasmania 1703

(a)

FIG. 7. Notoscolex pilus sp. nov., variations in genital markings: (a) P5; (b) P13.

P5-7, PII--12) and 20/21 (H, P2, P8-10 and P13-14 although markings are more ventral in these latter two specimens) and 21/22 (PI).

Internal anatomy Septa: thin. Gizzard: muscular in 5 (but displaced to 6). Oesophagus: white, dilated and lamellate in 14 and 15 forming annular calciferous glands. Nephridia: avesiculate meroic, two per side in ca. band d lines, intertwined but separable with care in fore-body, (appearing as holonephridia in hind-body); not tufted anteriorly. Vascularization: hearts 10-12; supra-oesophageal vessel weak in 9-12. Spermathecae: two pairs in 8 and 9; elongate ampulla on short duct with long, digitiform divertic­ ulum ectally (iridescent). Male organs: holandric, iridescent testes in 10 and 11 in seminal mucus; racemose seminal vesicles in 9 and 12. Ovaries: large in 13; ovisacs absent. Prostates: tubuloracemose (or flattened racemose?) in 18-20,21; duct external 1704 R. J. Blakemore and T. J. Kingston to the glandular part for most of its length (H and PI) and (in P 1 only) bilobed; long, curving penial setae sheathed in 18-23. Intestine: from 17 but septum 16/17 often displaced forwards; intestinal gizzards absent; typhlosole absent but low dorsal ridge sometimes present; gut contains gritty, yellow soil.

Remarks Notwithstanding the possibility that only species with multiloculate spermathecal diverticula (and extramural calciferous glands) are truly congeneric with the type­ species of Notoscolex, the present species is tentatively placed in this genus. Notoscolex species known from the north-west region that possess two pairs of spermathecae are N. simsoni, N irregularis (Spencer, 1895) and N bidiverticulatus (Jamieson, 1974), all differ from N pilus (based on reinspection of types and on new material), not least in the distribution of genital markings. Similar quadrithecal species from south-eastern Tasmania are N campestris (Spencer, 1895), N leai Michaelsen, 1910 and N wellingtonensis (Spencer, 1895), however these species have multiloculate spermathecal diverticula (and often a deep typhlosole). Some variations in the arrangements of genital markings appear in different populations of N. pilus as described above (cf. those from Wombat Hill and Pelion Valley). Specimens from these populations nevertheless conform on all other mor­ phological characters. The shape of the spermathecae, prostates and penial setae in Notoscolex pilus are reminiscent of Nexogaster sexies; this latter species differs in the distribution of its anterior genital markings, larger size, racemose prostates, in having a deep typhlosole and, most significantly, in having intestinal gizzards. Notoscolex pilus may represent a species having common descent with Nexogaster sexies from a precursor that was perhaps not unlike Megascolides maestus (below).

Distribution and habitat From Dismal Swamp Nature Reserve, Waratah and Pelion Valley in north-west/ central Tasmania.

Megascolides maestus sp. nov. (Fig. 8)

Material examined HOLOTYPE: (H) QVM:14:3327, Dismal Swamp Nature Reserve, N.W. Tasmania, 40 0 59'S 144°51'E, 8 September 1987, T. J. Kingston (complete mature, dissected and figured). PARATYPES: (PI) ANIC:RB.96.12.1, Dismal Swamp, approximately 400m along eastern reserve track, 3 December 1996, R. J. Blakemore and T. J. Kingston (com­ plete mature, dissected); (P2) TM:KI531, same details as PI (mature in two halves, dissected) . (P3) QVM: 14:3570, Dismal Swamp, gully west of turn off beside Bass Highway, 40°58'8 144°51 'E, 3 December 1996, R. J. Blakemore and T. J. Kingston (mature, caudal tip missing).

External features Lengths mm: ca 35. Width: 1·6mm. Segments: (H) 99, (PI) 105, (P2) 102. Colour: unpigmented, small yellow dots near setae c on anterior segments; clitellum New genus Nexogaster from Tasmania 1705

(c)

",-".._-.-_. abc d

(a)

FIG. 8. Megascolides maestus sp. nov., holotype: (a) tubular prostate and penial setae; (b) spermathecae; (c) prostomium. 1706 R. 1. Blakemore and T. J. Kingston pale yellow. Prostomium: open epilobous. Clitellum: }13-}17,17. Dorsal pores: small in 4/5, larger from 5/6. Nephropores: not found. Setae: eight throughout in regular series. Spermathecal pores: 7/8-8/9 in b lines. Female pores: paired in 14. Male pores: superficial in ab on 18. Genital markings: paired discs anterior to a setae on 10; pairs of discs in ab obscuring median intersegmental furrow in 17/18, and slightly narrower in 19/20-21/22 (three pairs in H; the last pair not developed in PI; only the first pair in P2-3 plus analogue on right in 20/21 in P2).

Internal anatomy Septa: thin. Gizzard: compact muscular in 5 (displaced to appear in 6). Oesophagus: narrow except for white dilated annulations (calciferous glands?) in 14 and 15. Nephridia: avesiculate meroic; two per side approximately in band d lines, intertwined and appearing as holonephridia but separable with care; not tufted. Vascularization: hearts 10-12; supra-oesophageal vessel seen in 9-12. Spermathecae: two pairs in 8 and 9; elongate saccular ampulla on thin duct with long and iridescent, digitiform diverticulum ectally. Male organs: holandric, iridescent testes in 10 and II in mucus; racemose seminal vesicles in 9 and 12. Ovaries: in '13; ovisacs absent. Prostates: tubular, coiled in 18-20, duct short; long penial setae in 18-23. Intestine: from 17 (septum 16/17 anteriorly displaced); gizzards absent although intestinal wall slightly thickened in 18-23; typhlosole absent; gut contains yellow soil, woody particles and some quartz grits.

Remarks Superficially Megascolides maeslus is similar to the sympatric NOlosco/ex pilus; major morphological differences are its plesiomorphic tubular prostates and, at the specific level, its characteristic markings in 10 and spermathecal pores in b, rather than a, setal lines. The occurrence of both these species in the same sample as specimens of Hickmaniella opisthogaster is of interest as this latter species may be derived from ancestral stock similar to M. maestus that has acquired tubuloracemose prostates (as in Notosco/ex pilus) and, additionally, perichaetine setae and an intest­ inal gizzard. Nexogaster sexies has undergone a different transition, attaining racem­ ose prostates while retaining lumbricine setae and deVeloping moniliform intestinal gizzards that must bestow an advantage, perhaps acting as 'grinding mills' using ingested quartz grits for more thorough comminution of its food. It is perhaps also significant that both M. maestus and N. pilus have calciferous gland-like modifica­ tions of the oesophagus whereas neither Hickmaniella spp. nor Nexogaster sexies has. The only previously described Megascolides from Tasmania is Megascolides acanthodriloides (Jamieson, 1974) which, amongst other differences, lacks calciferous glands. The distinctive male genital field of M. acanthodriloides was stated by Jamieson (1974) to distinguish this species from M. diaphanus from Victoria but also to demonstrate a 'special relationship' with Notosco/ex bidiverticulatus from Tasmania with which it has an almost identical genital field (as does Woodwardiella acanthodriloides Jamieson, 1971, figs. 2(a) and 4(b». The occurrence of Megascolides in Victoria, New South Wales and Western Australia (see generic definition above) serves to link the Tasmanian fauna with those of both the eastern and the south-western subregions of Australia. New genus Nexogaster from Tasmania 1707

Distribution and habitat Dismal Swamp Nature Reserve, sympatric with Hickmaniella opisthogaster, Anisochaeta simpsonorum and Notoscolex pilus and some other new endemic species, currently in the process of being described, as well as exotic lumbricids.

Acknowledgements A comprehensive two-year survey of native earthworms of Tasmania was under­ taken by the second author with funds made available by the Commonwealth of Australia under the National Estate Grants Program. The considerable earthworm collection accummulated under this grant has provided an ideal opportunity for the renewal of taxonomic studies for the region. The quality of the collection is largely due to the diligence of the National Estate Project Officer, Rob D'Orazio, in both the field and laboratory. Assistance in field provided by volunteer Daniel Soccol is also acknowledged with gratitude. Mr Stephen Pilkington is especially thanked for his cooperation with earthworm collection on his property 'Killara' near Marrawah. Mr R. Buttermore, Curator ofInvertebrates, Tasmanian Museum, Hobart is thanked for arranging the loan of type material from that institution. Taxonomic studies by the principal author while based at the Queen Victoria Museum were funded by a grant from the Plomley Foundation; the support of the trustees is greatly appreciated.

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